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"•I" 


BULLETIN  OF 

THE  BRITISH  MUSEUM 

(NATURAL  HISTORY) 


ZOOLOGY 

Vol.  26 

1973-1974 


BRITISH  MUSEUM  (NATURAL  HISTORY) 
LONDON:  1975 


DATES   OF  PUBLICATION   OF  THE   PARTS 

No.  i  .          .          .          .1  November  1973 

No.  2  .          .          .          .     15  November  1973 

No.  3  .          .          .          .          .        7  February  1974 

No.  4 22  May  1974 

No.  5 24  May  1974 

No.  6 7  June  1974 


ISSN  0007-1498 


Printed  in  Great  Britain  by  John  Wright  and  Sons  Ltd.  at  The  Stoncbridge  Press,  Bristol  BS4  5NU 


CONTENTS 

ZOOLOGY    VOLUME    26 

PAGE 

No.  i.  A  revision  of  the  large  Barbus  (Pisces,  Cyprinidae)  of  east  and 
central  Africa.  Studies  on  African  Cyprinidae  Part  II.  By 
K.  E.  BANISTER  .........  i 

No.  2.     Some  digenetic  trematodes  in  fishes  from  the  Bay  of  Biscay  and 

nearby  waters.     By  R.  A.  BRAY      ......  149 

No.  3.     Observations  on  Trinema  linear e  Penard  (Testacea  :  Protozoa). 

By  R.  H.  HEDLEY  and  C.  G.  OGDEN  (7  Pis.)     ....  185 

No.  4.     The   indigenous    earthworms    (Megascolecidae  :  Oligochaeta)    of 

Tasmania.     By  B.  G.  M.  JAMIESON  (10  Pis.)     ....  201 

No.  5.  The  freshwater  fishes  of  Rivers  Mungo  and  Meme  and  Lakes 
Kotto,  Mboandong  and  Soden,  West  Cameroon.  By  E.  TRE- 
WAVAS  (5  Pis.) 329 

No.  6.     Notes  on  some  echinoderms  from  southern  Africa.     By  A.  M. 

CLARK  (3  Pis.)       .........  421 

Index  to  Volume  26       ........  489 


INDEX  TO  VOLUME  26 


The  page  numbers  of  the  principal  references  and  the  new  taxonomic  names  are  printed  in  bold  type. 


abbreviatus,  Procatopus    .          .      377,  381,  384 
aberrans,  Procatopus         .        377,  379,  381,  384 
abyssorum,  Deretrema     ....      167 

abyssorum,  Proctophantastes  .         169-170,  172 
abyssorum  Steganoderma          151,  154,  167-ijo 
Acanthamoeba        .          .          .          .          .194 

acanthifer  phragmorus,  Astropecten  .      433 

Acanthodrilinae      .          .  203,  212-216,  263 

Acanthodrilini         .          .          .         203,  212-216 
acanthodriloides,  Pseudocryptodrilus  204,  214, 
296,  297,  298-299,  324,  Pis.  61-63 
Acanthophiothrix   .          .          .         465-466,  467 
acinus,  Trinema      .          .          .          .          .189 

aculeatus,  Prosorhynchus          .          .  159 

acuminatus,  Psilaster       ....      434 

acuticeps,  Barbus       3,  6-9,  27,  73,  89,  no,  128, 

140,  144 

acutirostris,  Barbus  brunellii    .          .  49,  60 

acutisquama,  Amphiura  .          .          .      423,  458 
adjecta,  Amphiura  .          .         446,  447,  483 

adustus,  Marcusenius       .          .          .       336,  338 
aeglefinus,  Gadus    .          .          .          .          .169 

affinis,  Barbus         .....        48 

afnnis  brevibarbis,  Barbus        ...        48 
affinis  nedgia,  Barbus      .          .          .          .48 

africana,  Asterias    .....     439 

africana,  Marthasterias    ....      439 

africana,  Marthasterias  glacialis         .          .     439 
africanus,  Asteracanthion          .          .          .      439 
Agrammobarbus     .....        94 

ahli,  Aphyosemion  .          .          .          .          -375 

ahli,  Auchenoglanis          .          .         331,  356-358 
ahlselli,  Barbus       ....  94-96 

albella,  Amphiura  .....     444 

albertisi,  Cryptodrilus     204,  208,  269,  270-271 
albertisi,  Megascolides      .          .          .       207,  270 
Alestes .          .          .          .  331,  339-340,  410 

alexandri,  Ophidiaster     ....      436 

alluaudi,  Astatoreochromis       .          .          .127 
alluaudi,  Barbus     .          .          .          .       3,  9,  144 

Alosa    ......       154,  178 

alsophila,  Perichaeta        .          .          .          .222 

altera,  Ophiopallas  paradoxa    .          .       477,  478 

altianalis,  Barbus        3,  7-8,  9-2O,  22,  25-27,  65, 

70,  73-74,  89,  94,  103,  106,  no, 

114,  127-128,  129-131,  134,  140,  144 

altianalis,  Barbus  altianalis          9-10,  12,  15-16, 

21,  23,  27,  144 
altianalis  altianalis,  Barbus  9-10,  12,  15-16, 

21,  23,  27,  144 

altianalis  eduardianus,  Barbus  9,  u,  15,  17-18, 
20,  21-23,  27»  XI4>  I27.  J44 


altianalis  labiosa,  Barbus          .          .  9,  103 

altianalis  lobogenysoides,  Barbus      .          .103 
altianalis  longifilis,  Barbus        .          .         76,  103 
altianalis  paucisquamata,  Barbus      .         76,  103 
altianalis  radcliffii,  Barbus  9-11,  13,  15,  17, 

19-20,  22-27,  I27.  I29.  X44 

altianalis  urundensis,  Barbus   .          .       in,  113 
alticola,  Barbus       .....        49 

altidorsalis,  Barbus  .          .          .          .119 

Amphichilus  .          .          .          .          .          .461 

Amphilepis     .....       448,  464 

Amphilimna  .          .          .  442—444,  479,  483 

Amphiodia     .          .          .          .          .       461,  481 

Amphiophiura         ....       423,  476 

Amphioplus       450,  451,  452-461,  463,  464,  481, 

483 

Amphipholis  .          .          .  450-452,  453,  461 

Amphispina  .          .          .          .          .       461,  481 

Amphiura   423,  444-450,  452,  454,  458,  475,  483 
Amphiuridae.          ....        444-464 

anacanthus,  Astropecten  .          .          .      432 

Anamphiura  ....         423,  478-479 

Andreasenius      376,  377,  379,  380,  381,  387,  413 
angolensis,  Ophiacantha  ....      442 

angularis,  Amphiura        .          .          .       446,  447 
angulosa,  Stereomyxa      .          .          .          .194 

annectens,  Labeo    ....       340,  341 

Anseropoda    ......      423 

Antedon         .....       427,  429 

Anthosticte    .          .          .          .          .          .423 

antinorii,  Aplocheilichthys        ...        72 
Antipodrilus  ......      205 

aper,  Capros  ....       154,  173 

Aphyosemion      331,  332,  370-376,  411,  412,  413 
apiocystis,  Diporochaeta  .          .          .223 

apleurogramma,  Barbus  .          .          .          .94 

Aplocheilichthys     .          .          .          .          .72 

Aplocheilus    ......      369 

appendiculatus,  Hemiurus        .          .       154,  178 
arabicus,  Barbus     ....        128,  131 

aranea,  Ophiopsila  .          .          .          .472 

arenilega,  Myxotheca       .          .          .          .195 

Argilophilus   .          .          .          .          .       218,  263 

Argulus  .  .  -35 

aristulata,  Ophiothrix      .          .         466-467,  468 
aristulata,  Placophiothrix         .          .       466,  467 
arnoldi,  Diporochaeta      .          .          .          .222 

Astatoreochromis    .          .          .          .          .127 

Asteracanthion        .....      439 

Asterias          ....         439,  440,  467 

Asteriidae       .....        439-440 

Asterina         ....         437~438.  483 


4QO 


INDEX 


Asterinidae    .  .        437-438 

Asterodiscides      .....     435 

Asterodiscus  ......      435 

Asteroidea      .....        431-440 

Astrocladus    ....       440-441,  PL  3 

Astroconus     .          .          .          .          .          .441 

Astropecten  .          .    423,  431-434,  483,  Pis.  1-2 
Astropectinidae       ....        431-434 

athertonensis,  Diporochaeta     .          .          .221 
athertonensis,  Perionyx  .          .          .          .221 

athi,  Barbus  .....  94-96 

Atopochilus 358,  366 

Auchenoglanis         .    331,  332,  356-358,  412,  4*3 
australis,  Barbus  intermedius          4,  50,  70-71, 

74,  75,  144 

avesiculatus,  Cryptodrilus     204,  208,  268,  269, 

270,  271-273,  275,  323,  Pis.  2-3 

avicularia,  Luidia   .....     431 

Azygiidae       .  .  .  .  .  .167 


babaulti,  Agrammobarbus 
babaulti,  Barbus     . 
babaulti,  Varicorhinus     . 
baccata,  Ophiacantha 
Bagridae 

bakeri,  Perichaeta  . 
ballayi,  Auchenoglanis     . 
Banister,  K.  E. 
Barbodes 


94 

4.  94-96 
.        94 

•  442 
,  356-358 

.      223 

•  357 
1-148 

•  349 


Barbus    1-148,  331,  332,  333,  341-343,  344-356, 

410,  411,  412,  413 
barbus,  Barbus        .          .          .          .          .134 

Barilius  ......        25 

barronensis,  Perichaeta   .          .          .          .222 

bassana,  Perionychella     .  203,  206,  253-255 

bassanus,  Megascolides    .          .         205,  ,222,  253 
bassanus,  Plutellus  ....      253 

batesii,  Barbus  128,  331,  342,  343-345,  346, 

412,  413 

batesii,  Chiloglanis  .  358,  359,  361,  364 

batesii,  Chromidotilapia  332,  392,  394,  395, 

396,  397,  401 
batesii,  Labeo          ....       340,  341 

batesii,  Marcusenius         ....      338 

batesii,  Pelmatochromis  .          .          .       393,  394 
Bathybiaster  .....     434 

Bathycreadium       .          .  151,  154,  160-162 

baudoni,  Barbus     ....       332,  356 

bayoni,  Barbus        ....  10-11 

beauforti,  Ophioteresis     .          .          .       469,  483 
beauforti,  Ophiothela       .          .          .       469,  483 
Beirabarbus  ......      347 

bemini,  Tilapia        ....       405,  411 

Benthotrema  .          .          .          .          .178 

Beryx   .  ...       154,  170 

bidiverticulatus,  Oreoscolex         204,  210,  298, 

303,  304,  305-307,  324,  PL  67 

bimaculatus,  Hemichromis       .          .       411,  413 

bingeri,  Barbus       .          .          .          .  49,  56 


bingeri,  Capoeta  .  .  .  .  .49 
bingeri,  Varicorhinus  ....  49 
biscayense,  Bathycreadium  151,  154,  160-I&2 
bispinosa,  Ophiopsila  .  .  472-475,  483 
bithecata,  Perionychella  203,  206,  221,  225, 

230,  232,  233-234,  256,  PL  93 
bithecata,  Vesiculodrilus  203,  206,  221,  225, 

230,  232,  233-234,  256,  PL  93 
bivittatum,  Aphyosemion  331,  370-371,  411, 

4J2,  413 

bivittatus,  Fundulus  .  .  .  .370 
blackeri,  Steringophorus  151,  154,  174-178 

blennoides,  Phycis  .  .  154,  162,  180-181 

bottegi,  Barbus  .....  49 
boulengeri,  Chromidotilapia  .  .  .401 
Brachyalestes  .  .  .  .  -339 
brachyistius,  Brienomyrus  331,  332,  336-338, 

410,  413,  PL  2 
brachyistius,  Marcusenius  .  335,  336,  338 

Bray,  R.  A 149-1 83 

bredoi,  Varicorhinus         .          .          119-120,  123 
brevibarbis,  Barbus          ....        48 

brevibarbis,  Barbus  affinis        ...        48 
brevibarbis,  Chiloglanis  ....      364 

brevispinis,  Barbus  .          .          .       342,  347 

Brienomyrus     331,  332,  335-338,  41°,  4X3,  PI-  2 
Brissidae        ......      480 

brucii,  Barbus         .          .          .          .          -37 

brucii,  Varicorhinus  .  .  .  37,  123 
brunellii,  Barbus  .  .  49,  60,  65,  69-70 

brunellii  acutirostris,  Barbus  .  .  49,  60 

brunyensis,  Cryptodrilus  204,  208,  268,  269, 
270,  273-275,  323,  Pis.  4-7 

Brycinus 339 

Bucephalidae  ....  155-159 

Bucephalopsis  ....  154,  157 
burniensis,  Megascolex  204,  214,  319-321,  324 
burtoni,  Asterina  ....  437-438 
burtonii,  Asterina  .  .  .  .  -437 
bynni,  Barbus  3,  20,  25-26,  27-3&,  45-47,  50, 
74,  78,  100,  114,  127-128,  140,  144 
bynni,  Cyprinus  ....  27-28 

bynni  complex,  Barbus  .  128-129,  131-133. 

139-140 

bynni  rudolfianus,  Barbus  .  .  27-28 

byrni,  Barbus  .....  33 


cabrae,  Tilapia  .....  406 
caelorhinchus,  Coelorinchus  .  154,  160,  167 
caeruleus,  Aphyosemion  calliurus  .  -375 
caeruleus,  Megascolex  .  .  .  -319 
caeruleus,  Panchax  calliurus  .  .  -375 
calabaricus,  Calamoichthys  .  .  .411 
Calamoichthys  .  .  .  .  .411 
callichaeta,  Woodwardia  .  .  .  265 

callipterus,  Barbus    331,  333,  347-349,  352,  411, 

413 

callipterus,,  Enteromius  .          .  347-349 

calliurum,  Aphyosemion .          .         373,  375,  376 


INDEX 


491 


calliurus  caeruleus,  Aphyosemion  .  .  375 
calliurus  caeruleus,  Panchax  .  .  .  375 
Callochromis .  .  .  .  .  39 1 

cameronense,  Aphyosemion  .  .  375,  376 
cameronensis,  Aphyosemion  .  .  373,  375 
cameron ensis,  Panchax  .  .  .  -375 
cameronensis,  Tilapia  ....  403 
camerunensis,  Labeo  331,  340-341,  413,  PI.  3 
camerunensis,  Tilapia  .  33 1,  332,  402-403, 

405,  411,  413 

camerunensis,  Tilapia  lata  .  .  .  402 
campestris,  Cryptodrilus .  .  .  205,  307 
campestris,  Notoscolex  ....  307 
campestris,  Oreoscolex  .  204,  210,  303,  305, 
306,  307-309,  324,  Pis.  68-72 
camptacanthus,  Barbodes  .  .  .  349 
camptacanthus,  Barbus  .  331,  349-35O,  352, 

354.  355.  4J3 
camptacanthus,  Puntius ....      349 

canaliculata,  Perichaeta  .          .          .       217,  221 
canaliculata,  Perionychella       .          .          .217 
canaliculatus,  Perionychella      .          .          .216 
Candida,  Amphiura  .          .          .       445,  483 

canis,  Barbus  .          .          .          .          .134 

canis,  Tor       ......      134 

capensis,  Amphiura          .          .         446-447,  483 
capensis,  Astroconus        .          .          .          .441 

capensis,  Barbus     .  .      .          .          .129 

capensis,  Dipsacaster  sladeni    .          .          .     434 
capensis,  Hacelia    ....       435-436 

capensis,  Hacelia  superba          .          .       435,  483 
capensis,  Ophiarachnella  .          .          .     480 

capensis,  Ophiogymna     ....      423 

capensis,  Ophiomyxa  vivipara.  .  .482 
capensis,  Perionychella  203,  206,  220,  223, 

224,  225-226,  228,  229,  256, 

258,  260,  Pis.  91-92 

Capoeta          .          .          .          .          .  49,  94 

Capros  ......       154,  173 

carnea,  Ophiactis    .....      465 

carpio,  Cyprinus     .          .          .          .          .128 

castellanii,  Acanthamoeba  .  .  .194 
caudifasciatus,  Chromidotilapia  .  .  395 
caudovittatus,  Barbus  3,  n,  36-41,  95,  106, 

122-123,  126,  130,  139,  144 

celiae,  Aphyosemion  331,  371-373,  412,  413 

Ceramaster    ....         423,  435,  483 

chadwicki,  Decametra     ....     426 

Channa  .          .          .          .          .          .411 

chapini,  Varicorhinus       .          .          .       123,  126 
Chara    .......      194 

Characidae     .  331,  339-34° 

chariensis,  Labeo    .          .          .          .          .341 

chariensis  nunensis,  Labeo        .          .          .341 
Charitometridae      ....        429-430 

Chiloglanis     .      331,  332,  358-361,  362-367,  413 
chilotes,  Barbus      .          .          .          .          .119 

chilotes  sakaniae,  Barbus          ...        36 
chlorotaenia,  Barbus        ....      354 

chondriscus,  Ceramaster .  .  .  435,  483 


chondriscus,  Ceramaster  patagonicus          .     435 

Chromichthys          ....       387,  388 

Chromidotilapia      .    331,  332,  888-393,  394-402, 

404,  411,  413,  Pis.  4-5 

chuni,  Democrinus.  ....  482 
Cichlidae  ....  331,  387-410 
cinereus,  Muraenesox  .  .  .  .157 
cinnamomeum,  Aphyosemion  .  .  -372 
Clarias  .  .  .  331,  356,  411,  412,  413 

Clariidae         .          .          .          .  331,  356 

Clark,  A.  M 421-487,  3  Pis. 

clarki,  Odinia          .....     482 

codringtonii,  Barbus         .          .          .          .119 

coelorhynchi,  Lepidapedon       .          .          .166 
Coelorinchus  ....         154,  160,  167 

Coitocaecum  .          .          .          .          .          .162 

Colobometridae       ....        424-426 

Comanthus     .....       480,  481 

comata,  Ophiothrix          ....      465 

compinei,  Barbus    .....      344 

complanatum,  Trinema  .          .          .          .189 

complanatum  globulosa,  Trinema      .          .189 
compressa,  Amphiura      .          .         446,  447,  483 
Conger .  .    154-155,  157-158,  178,  181 

conger,  Conger        .    154-155,  157-158,  178,  181 
congicus,  Chiloglanis        .          .          .       365,  366 
coniortodes,  Amphioplus  .  .  .      464 

copei,  Xenodermichthys  .          154,  174-176 

copelandi,  Perichaeta       .          .          .          .220 

Coptodon        ......      403 

corynephora,  Ophiomitrella      .          .       441,  482 
Coryphaenoides       .          .          .          .          .169 

cribriformis,  Amphilimna      .         442-444,  483 
Crinoidea        .....        424-430 

Crotalometra  .          .  427-429,  430,  482 

crucibulum,  Distoma       .          .          .          .157 

crucibulum,  Gasterostomum     .          .          .      157 

crucibulum,  Monostoma  .          .  157 

crucibulum,  Prosorhynchus       151,  154,  157-159 

Cryptodrilus      204,  205,  208,  212,  214,  219,  221, 

222,  241,  247,  257,  260,  265,  266-296, 

3°3.  3°7,  309.  317.  323,  Pis-  2-59 

Ctenopoma    .          .          .          .          .411,  413 

Curimatus      ......      389 

curtus,  Barbus        .          .          .          •          .116 
cylindricus,  Labeo  .          .          .          .          .341 

Cyprinidae     .  .  .       1-148,  331,  340-356 

Cyprinodontidae     .          .          .         331,  367~387 
cyprinoides,  Mormyrus    ....      335 

Cyprinus 27-28,  128 


daga,  Barbus  platystomus         ...        48 
dageti,  Tilapia         .          .  .      403 

dainellii,  Barbus     .          .          .          -49.  52,  55 
dainellii  macrocephalus,  Barbus         .          .        49 

Dalatias 154 

danae,  Ophiothela  .....     470 

dartnalli,  Cryptodrilus  204,  208,  269,  270, 

273,  275-277,  323,  Pis.  8-12 


492 


INDEX 


davallia,  Diporochaeta    ....     220 

davallia,  Perionychella    .          .          .          .217 

davidis,  Antipodrilus       .  .          .     205 

decadactylus,  Beryx         .          .          .       154,  170 
Decametra     ....        423,  425,  426 

decatheca,  Perionychella  .        203,  206,  255 

decatheca,  Plutellus         .          .        207,  222,  255 
degeni,  Barbus        .          .          .         49~5°>  54~55 
degeni  leptorhinus,  Barbus       ...       49 
dekkensis,  Barbus  platystomus          .          .       48 
Democrinus   ......      482 

dendyi,  Perichaeta.          .          .          .       217,  218 

dendyi,  Perionychella      .          .          .        217  220 
Deretrema     .....       167,  169 

Dermechinus.          ....        479-480 

Derogenes      ....         154,  180-181 

deserti,  Barbus        .....      348 

diaphanus,  Megascolides ....     296 

diaphanus,  Pseudocryptodrilus          .       296,  299 
Dichogastrini          .          .          .         204,  300-302 
dicksonia,  Perichaeta       ....     223 

Difflugiella     .          .          .          .          .          .196 

Digenea          .....        149-183 

dilwynnia,  Diporochaeta  .  .  .  234 

dilwynnia,  Perichaeta  .  .  205,  221,  234 
dilwynnia,  Perionychella  203,  206,  225,  232, 

234-236,  256,  Pis.  94-96 
dilwynnia,  Vesiculodrilus  203,  206,  225,  232, 

234-236,  256,  Pis.  94-96 

Diporochaeta    204,  217,  218,  219,  220,  221,  222, 

223,  228,  234,  245,  257,  258,  259 

Dipsacaster    ......     434 

disneyi,  Chiloglanis         .         331,  361,  362,  363, 

355-367,  413 
Distoma         .....       157,  173 

Distomum      .          .          .          .          .          .162 

divergens,  Distoma  .          .          .          .173 

divergens,  Fasciola  .          .          .   •       .      173 

divergens,  Steringotrema  151,  154,  173-174 

dividua,  Amphiodia         .          .          .          .461 

dividua,  Ophiothela         ....      470 

Dolichoenterum      .          .  151,  154,  155-157 

donyensis,  Barbus  ....  94~95 

dubia,  Perichaeta  .          .          .          .          .220 

dubia,  Tilapia          .....      405 

dubia,  Tilapia  mariae      .          .          .       405,  406 
dubius,  Cryptodrilus        .          .          .       269,  282 
dubius,  Microscolex          .          .          .          .213 

duchesnii,  Barbus  .....        48 

duchesnii  ibridus,  Barbus          ...        48 
duchesnii  maximus,  Barbus      ...        48 
durbanensis,  Decametra  .          .        423,  425,  426 
dyscrita,  Asterina  ....       438,  483 

dyscrita,  Patiriella.  ....  438 


eburneensis,  Barbus  .  .  .  354,  355 
Echinaster  ....  438,  439,  483 
Echinasteridae  ....  438-439 
echinata,  Ophiothrix  fragilis  .  .  .  468 


Echinidae 
Echinodermata 
Echinoidea     . 
Echinus 
eduardianus,  Barbus 


479-480 

.     421-487,  3  Pis. 
479-480 

•      479 
.   9-n 


eduardianus,  Barbus  altianalis  9,  n,  15,  17-18, 
20,  21-23,  27,  IJ4.  I27,  144 
electricus,  Malapterurus  .  .  331,  367,  413 
electricus,  Silurus  .....  367 
elegans,  Asterodiscides  .  .  .  435 

elegans,  Asterodiscus  ....  435 
ellisi,  Cryptodrilus .  .  .  205,  222,  257 
ellisi,  Perionychella  .  .  203,  206,  257 

ellisi,  Plutellus  .  .  .  .  .257 
elongata,  Nicolla  .  .  .  .  .162 
elongatum,  Bathycreadium  .  .  151,  162 
elongatum,  Lepidapedon  151,  154,  164-1 66 

elongatum,  Lepodora  .  .  .  .164 
elongatum,  Nicolla  .  .  .  .151 

elongatum,  Steganoderma  .  .  .169 
elongatus,  Barbus  .....  48 
elongatus,  Chromichthys  .  .  387,  388 

elongatus,  Hemichromis  .          331,  332,  387-388, 

411.  413 

enchelys,  Trinema  ....        188-189 

Enteromius    .....        347-349 

enteronephricus,  Cryptodrilus      204,  208,  267, 

268,  269,  273,  275,  276,  277-28o, 

286,  290,  323,  Pis.  13-19 

Epiplatys       .  .  331,  367-370,  411,  413 

eques,  Lepidion       .          .  154,  160,  164,  167 

erici,  Diporochaeta  .          .          .          .221 

erici,  Perionyx  .  .  .  .  .221 
erlangeri,  Barbus  .  .  .  49,  55,  57,  72 
esculenta,  Tilapia  .  .  .  .  .391 
esculentus,  Sarotherodon  .  .  -391 

ethiopicus,  Barbus.          .     3,  41-44,  9O-9X.  129, 

139,  144 
euchilus,  Barbus     .....        36 

Euglypha       .          .          .  187-188,  192,  195-197 
Euglyphacea.          .          .          .          .          .187 

Euglyphidae  .          .          .          .          .          .188 

eumystus,  Barbus  .....        49 

Euryale  ......      440 

euryale,  Asterias     .....      440 

euryale,  Astrocladus         .          .       440-441,  PL  3 
euryplax,  Ceramaster  patagonicus     .       435,  483 
euzona,  Diporochaeta      .          .          .          .222 

evansi,  Perionychella  203,  206,  221,  225,  230, 

235,  236-237,  254,  256,  258 
evansi,  Vesiculodrilus  203,  206,  221,  225,  230, 

235,  236-237,  254,  256,  258 

excisum,  Lecithocladium  .  .  154,  178 

exigua,  Asterina  .....  438 
exigua,  Patiriella  .....  438 


fairbairnii,  Barbus . 
falcatus,  Amphioplus 
falcatus,  Unioplus  . 


.      119 

•  455-456,  458,  459,  460 
455-456 


INDEX 


493 


fallax,  Alosa 154,  178 

fasciata,  Helicometra       .          .          .       154,  159 
fasciatus,  Hemichromis     332,  387,  388,  411,  413 

Fasciola 173 

fastigatus,  Cryptodrilus  .          .         269,  282,  288 
faucium,  Diporochaeta    ....      223 

Fellodistomidae       ....        173-178 

Fellodistomum        .          .          .          .          .177 

fergussonii,  Barbus  ....   9-11 

Filosia  .......      187 

finleyi,  Chromidotilapia .          331,  392,  393-397, 

400,  411,  413,  PI.  5 
finta,  Alosa    .          .          .          .          .          .178 

Fletcherodrilus        .          .          .          .       218,  221 

flexicollis,  Bathycreadium         .          .          .162 
flexuosa,  Ophiactis  .          .          .       464,  465 

floridae,  Hacelia      .....     436 

formosum,  Steganoderma          .          .          .      169 
forskalii,  Labeo       .          .          .          .          .341 

foureaui,  Barbus     .          .          .         344,  346,  347 
fragilis,  Asterias      .....     467 

fragilis,  Ophiothrix  .          .         467-469,  483 

fragilis  echinata,  Ophiothrix     .          .          .      468 
fragilis  lusitanica,  Ophiothrix  .          .          .      468 
fragilis  pentaphylla,  Ophiothrix         .          .      468 
fragilis  pentaphyllum,  Ophiothrix     .          .      468 
frenchi,  Cryptodrilus        .          .          .       221,  222 
frenchi,  Megascolex          ....      326 

frenchi,  Vesiculodrilus      .          .          .          .222 

frosti,  Diporochaeta         .          .          .          .222 

Fundulus        .          .          .          .          .          -370 

furcatus,  Amphioplus         452-453,  454,  481,  483 
furcatus,  Lymanella         .  452-453,  481,  483 


gadi,  Lepidapedon  .          .          .          .166 

Gadus   ......       166,  169 

Gaidropsarus  ....       154,  159 

galeata,  Trinema     .          .          .          .          .189 

galilaea,  Tilapia      .....      408 

galilaeus,  Sarotherodon  .  .  331,  332,  404, 

408-410,  412,  413 

galilaeus,  Sarotherodon  galilaeus  .  409,  411 
galilaeus,  Sparus  .....  408 
galilaeus  galilaeus,  Sarotherodon  .  409,  411 
galilaeus  multifasciatus,  Sarotherodon  .  408 
gananensis,  Barbus  .  3,  32,  36,  44-47,  ?8, 

100,  128,  140,  144 
Garra    .....  90,  131,  389 

Gasterostomum       .          .          .          .  157 

Geophagus     .          .          .          .          .          .391 

georgei,  Graliophilus        ....      260 

gibbosus,  Amphioplus      .          .         452,  453,  454 
gibbosus,  Ophiophragmus         .         452,  453,  454 
gigas,  Mithrodia      .          .          .          .          .423 

gippslandicus,  Cryptodrilus       .          .          .      222 
glacialis,  Marthasterias    ....      439 

glacialis  africana,  Marthasterias  .  .  439 
glacialis  rarispina,  Marthasterias  .  .  439 


glandifera,  Perionychella       203,  206,  218,  221, 

223,  230,  237-238,  254,  256 

glandifera,  Vesiculodrilus      203,  206,  218,  221, 

223,  230,  237-238,  254,  256 

glandulosa,  Proctophantastes  .          .          .170 
glandulosum,  Neosteganoderma         .      151,  154, 

170-173 

glaucicaudis,  Procatopus          331,  377,  379,  380, 
381,  382,  383,  384,  386 

globulosa,  Trinema  complanatum      .          .189 
globulosa,  Trinema  lineare        .          .          .189 
Glyptometra .          .          .  428,  429-430,  482 

Gnathonemus          .          .          .          .          -338 

goheeni,  Mormyrus  .          .         334,  335,  413 

Goniasteridae  .....      435 

Gorgonocephalidae  .          .          .        440-441 

Gorgonocephalus     .....      440 

gorgorensis,  Barbus  intermedius        .          .        48 
gorguarii,  Barbus    .          .    48,  55,  65,  68-69,  128 
gorguarii  macrophtalmus,  Barbus     .          .        48 
gracilescens,  Bucephalopsis       .          .       154,  157 
gracilis,  Procatopus          ....      380 

gracilispina,  Asterina       ....     437 

Graliophilus  .    203,  204,  209,  212,  214,  218,  219, 

259-26o,  261-262,  263,  265,  324,  PI.  i 

grandis,  Diporochaeta      .          .          .          .222 

grandis,  Notoscolex          ....      303 

grandis,  Perionychella     .          .          .          .216 

grandis,  Vesiculodrilus     .          .  .      .      216 

grandisquama  natalensis,  Amphiura         447-448 
granulatus,  Astropecten  .          .         431,  432,  433 
granulatus  natalensis,  Astropecten  423,  431-433, 

PI.  i 

gravidum,  Lecithochirium         .         154,  178-i8i 
greenii,  Labeo         .          .          .          .          .341 

gregorii,  Barbus    3,  4,  10,  48,  50,  70,  72,  75,  143 
Gromia  ......      195 

Gromida         ......      187 

gudaricus,  Barbus  .....        49 

guentheri,  Atopochilus     ....      358 

guineensis,  Tilapia  ....       402,  403 

guntheri,  Chromidotilapia        332,  389,  390,  391, 

392,  393,  394,  398-402,  411,  413 

guntheri,  Chromidotilapia  guntheri  .      331,  398, 

399,  400,  401 
guntheri,  Hemichromis    ....      398 

guntheri  guntheri,  Chromidotilapia  .      331,  398, 

399,  400,  401 
guntheri  loennbergi,  Chromidotilapia      331,  391, 

392,  398,  399,  4°°.  401.  4",  P1-  4 
gymnophallum,  Coitocaecum  .  .  .162 
Gymnophiura  .  .  .  475,  476,  483 


habereri,  Barbus     . 

Hacelia 

halli,  Megascolex     . 

hamata,  Ophiomitrella 

Haplochilus   . 

Haplochromis 


•       346,  347 
435-436,  483 

•  326 
.      482 

•  367 
26,  389,  391 


494 


INDEX 


harringtoni,  Barbus 
hastatus,  Amphioplus 
hastatus,  Lymanella 
haugi,  Pelmatochromis 
haugi,  Tilapia 
healesi,  Woodwardia 
Medley,  R.  H. 
Helicometra  . 
hellyeri,  Perionychella 


452,  481,  483 
.  481 
402 

405,  406 
219 

185-iQQ,  7  Pis. 
154.  159 

203,  2O6,  221,  225, 
230,  235,  288-241,  247,  256 

hellyeri,  Vesiculodrilus  .         203,  206,  221,  225, 

230,  235,  238-241,  247,  256 

Hemichromis      331,  332,  387-388,  398,  411,  413 


Hemiuridae    . 

Hemiurus 

Henricia 

heteroporus,  Plutellus 

heudeloti,  Tilapia   . 

hickmani,  Perionychella 

Hickmaniella 


178-181 
154,  178 
438-439,  483 
.  268 

•   4°5 

203,  206,  220,  223, 
226-228,  229,  256 

204,  2O5,  209,  212,  214,  270, 

300-302,  324,  Pis.  64-66 

hindii,  Barbus  ....  94-97,  127 
hobartensis,  Cryptodrilus  .  205,  221,  241 
hobartensis,  Perionychella  203,  206,  225,  230, 
241-245,  250,  251,  253,  254,  255,  256,  Pis.  97-98 
hobartensis,  Plutellus  .  .  .  .241 
hobartensis,  Vesiculodrilus  203,  206,  225,  230, 
241-245,  250,  251,  253,  254,  256,  Pis.  97-98 
hoggi,  Megascolex  .....  326 
hollyi,  Barbus  .  .  .  .  .10-12 
holubi,  Barbus  .  .  .  .  .129 
horridus,  Dermechinus  .  .  .  479~48o 
horridus,  Echinus  .....  479 
huguenini,  Barbus  ....  83 

humilis,  Barbus  .  .  .  .  .114 
hursensis,  Barbus  ....  49,  55-56 
Hydrocynus  .  .  .  .  .  '  .  26 

Hylopanchax  .....      382 

Hypsopanchax        ....       377,  382 


ibridus,  Barbus  duchesnii          ...        48 
ilgi,  Barbus    ......        49 

illawarrae,  Cryptodrilus  ....      303 

imparicystis,  Oreoscolex  ....      303 

incana,  Amphiura  .....      452 

infrafasciatus,  Haplochilus        .          .          .      367 
insularis,  Cryptodrilus      .  .  .        205,  241 

insularis,  Perionychella    ....      255 

insularis,  Plutellus .          .          .          .          .241 

insularis,  Vesiculodrilus  .          .          .          .241 

integer,  Amphioplus  450,  451,  452,  453-455,  461 
integer,  Lymanella     450,  451,  452,  453-455,  461 
integra,  Amphipholis       .          .          .          -453 
integra,  Amphiura  .....      454 

intermedia,  Diporochaeta          .          .       219,  220 

intermedius,  Barbus  3,  14,  16,  27,  33,  44,  47-74, 

91,  no,  127-128,  130-132,  134,  140,  144 

intermedius,  Barbus  intermedius    71,  74-75,  144 


intermedius  australis,  Barbus 

intermedius  complex,  Barbus    . 

114,  128-129, 

intermedius  gorgorensis,  Barbus 
intermedius  group,  Barbus 
intermedius  intermedius,  Barbus 

intermedius  leptosoma,  Barbus 
intermedius  microstoma,  Barbus 
inutilis,  Ophiocirce 
irregularis,  Cryptodrilus  . 
irregularis,  Diporochaeta 
irregularis,  Notoscolex     . 
irregularis,  Oreoscolex 


irregularis,  Perichaeta 
irregularis,  Perionychella 


204, 
306 


227 


4.  50,  70-71, 
74,  75,  144 

73,  74-  "0, 

131-135,  140 

.        48 

•     9,  73-  129 

.    71,  74-75, 

144 

.        48 

.        48 

423,  476-477 

205,  309 

.      228 

•      309 

210,  303,  305, 
309-3IO,  324 
205,  22O,  228 

2O3,  2O6,  223, 

,  228-229,  256 


Jamieson,  B.  G.  M.  .  201-328,  102  Pis. 

japonica,  Polymixia  .  .  .  .172 
jarsinus,  Barbus  .....  48 
jobaerti,  Hypsopanchax  ....  382 
jubbi,  Barbus  .....  82 
jurupari,  Geophagus  .  .  .  .391 


kalki,  Amphiura  ....  445,  483 
kassamensis,  Barbus  .  .  .  48,  55-56 
kerguelensis,  Leptoptychaster .  .  .  434 
kershawi,  Perionychella  203,  206,  220,  223, 

224,  229-230,  256 

kerstenii,  Barbus  .  .  .  .  .129 
kimberleyensis,  Barbus  .  .  .  .129 
kingsleyae,  Chromidotilapia  .  388,  395,  402 
kingsleyae,  Ctenopoma  .  .  .411,  413 
kiogae,  Barbus  .  .  .  .  .10 
kivuensis,  Barbus  .  .  .  .  .10 
koreae,  Amphiura  .....  450 
kottae,  Tilapia  .  3 31,  332,  403,  404-405, 

407,  408,  409,  411,  412 
krapfi,  Barbus         ....     94-96,  98 


Labeo   .  135,  331,  332, 

Labeobarbus . 

labiatus,  Barbus 

labiosa,  Barbus  altianalis 

Labyrinthula 

lacustris,  Diporochaeta    . 

lacustris,  Perionychella    . 


lacustris,  Perionyx. 
lacustris,  Procatopus 

lacustris,  Vesiculodrilus 
lagensis,  Barbus 


225 


382 


340-341,  413,  PL  3 

.  9,  48 

•   94-98,  127 

9,  103 

•  194 

•  245 

2O3,  2O6,  221,  223, 

,231,  245-246,  256 

207,   221,  245 

331-  378,  379,  380, 
,  383,  386-387,  412 

203,  221,   223,  225, 

231,  245-246,   256 

83,   130,   132 


INDEX 


495 


lamottei,  Chiloglanis 
Lanceabarbus 
lancrenonensis,  Barbus    . 
lapsus,  Barbus 
lata,  Tilapia  . 
lata  camerunensis,  Tilapia 
Lates     .... 
leai,  Notoscolex 


365,  366,  367 
94 

•  347 

•  4-  94 

402 

402 

26 

207,  310 


leai,  Oreoscolex       .    204,  210,  303,  305,  310-3  n 
Lecithochirium        .          .          .          154,  178-181 
Lecithocladium       ....       154,  178 

Lepidapedon.          .          .  151,  154,  162-166 

Lepidion         .          .          .  154,  160,  164,  167 

lepidus,  Barbus       .....      355 

Lepocreadiidae        ....        162-166 

Lepodora        .....        162,  164 

Leptoptychaster      .....     434 

leptorhinus,  Barbus  degeni       ...        49 
leptosoma,  Barbus.          .          .          .48,  50,  no 

leptosoma,  Barbus  intermedius          .          .        48 
lestradei,  Barbus     ....     36-38,  95 

Lethrinops      .  .  .  .  .  .391 

liberiensis,  Barbus  .  .         332,  354,  355 

liberiensis,  Marcusenius   .          .         336,  337,  338 
licha,  Dalatias         .          .          .          .          .154 

limicola,  Ophionephthys .          .         462,  463,  464 
Limnodrilus  ......      205 

lindti,  Diporochaeta         ....      223 

lineare,  Trinema     .          .  188-199,  7  Pis. 

lineare  globulosa,  Trinema        .          .          .189 
lineare  pellucida,  Trinema         .          .          .189 
lineare  terricola,  Trinema          .          .          .189 
linearis,  Amphiura ....       423,  475 

linnellii,  Barbus      .....      344 

Lithodesium  ......      194 

littoralis,  Rhododrilus   .  203,  213-2x6,  297 

324,  PI.  60 

lobogenys,  Barbus  .          .          .  10,  14-15 

lobogenysoides,  Barbus  altianalis       .          .103 
lochensis,  Perichaeta        .          .          .          .222 

loennbergi,  Chromidotilapia      .          .       332,  392 
loennbergi,  Chromidotilapia  guntheri      331,  391, 
392,  398,  399,  400,  401,  411,  PI.  4 
longianalis,  Brienomyrus  332,  336,  337,  410 

longianalis,  Marcusenius .          .          .       336,  338 
longinlis,  Barbus      3,  4,  n,  36,  76-79,  100,  106, 

128,  140,  145 

longinlis,  Barbus  altianalis        .  .          76,  103 

longipinnis,  Alestes  331,  339-340,  410,  413 

longipinnis,  Brachyalestes         .          .          -339 
longirostris,  Barbus          .          .  10,  12,  16-17,  J9 
longispinis,  Barbus  ....        83 

longissimum,  Dolichoenterum   151,  154,  155—157 

longus,  Oreoscolex  204,  210,  303,  304,  305, 

311-313,  324,  Pis.  73-80 

Lophaster       .          .          .          .          .          .438 

Lophius          .....       154,  157 

lowei,  Polymixia     .  .  .  .  .172 

lowelli,  Ophionephthys  .          .         462-464,  483 
luetkeni,  Ophiactis  ....      465 


Luidia  .          .          .          .          .          .          .431 

Luidiidae        .  .  .  .  .  .431 

Lumbri'culus  ......      205 

lusitanica,  Ophiothrix  fragilis  .          .          .      468 
Lymanella  450,  451,  452-455,  461,  481,  483 

lymani,  Ophiactis   ....       464,  465 


maclareni,  Clarias  .....  356 
macmillani,  Barbus  .  .  -49,  55,  57 
macrobrachius,  Oligometra  serripinna  .  426 
macrocephala,  Tilapia  .  .  .  389,  408 
macrocephalus,  Barbus  dainellii  .  .  49 
macrolepidotus,  Alestes  .  .  331,  339,  413 
macrolepidotus,  Brycinus  .  .  -339 
macrolepis,  Barbus  3,  79-83,  130,  132,  139, 

145 

macronema,  Barbus         .          .          .     48,  68-69 
macronema  parenzani,  Barbus  .  48,  52 

macrophtalmus,  Barbus  gorguarii      .          .        48 
macrophthalma,  Molva    .          .          154,  180-181 
macropus,  Barbus  .....        83 

Macrurus        .          .          .          .          .          .169 

maculata,  Luidia    .          .          .          .          .431 

magnicirra,  Antedon        ....      427 

magnicirra,  Crotalometra  427-429,  430,  482 

magnifica,  Tropiometra   .          .         423,  426-427 
magnus,  Steringophorus  .          .          .          .178 

makiensis,  Garra     .....        90 

Malapteruridae        .          .          .          .       331,  367 

Malapterurus  .          .          .         331,  367,  413 

manni,  Diporochaeta        ....      222 

manteri,  Dolichoenterum  .  .  .157 

Marcusenius  .  335,  336,  337,  338 

marequensis,  Barbus         4,  10,  82,  119,  123,  130 
margaritae,  Barbus  ....        49 

mariae,  Barbus        .   3,  83-87,  103,  129,  139,  145 
mariae,  Tilapia        .  331,  332,  389,  404,  405-4o8, 

411,  412,  413,  PI.  4 

mariae  dubia,  Tilapia       .          .          .       405,  406 
marlieri,  Chiloglanis         .          .         365,  366,  367 
Marthasterias  .....      439 

mathoiae,  Barbus   ....     94-96,  98 

matris,  Barbus        .....        83 

mawambi,  Barbus  .....        94 

mawambiensis,  Barbus    .          .  95,  no-in,  344 
maximus,  Barbus  duchesnii      ...        48 
mediocincta,  Diporochaeta        .          .          .      222 
mediocris,  Cryptodrilus   .          .          .       266,  269 
meeki,  Tilapia         .....      405 

Megascolecidae        .          .  201-328,  102  Pis. 

Megascolecinae        .          .          .         203,  216-326 
Megascolecini  .          .  204,  280,  302-326 

Megascolex     .          .  204,  205,  212,  214,  217, 

220,  318-326 

Megascolides  .     205,  207,  222,  253,  263,  270,  296 
melanopleura,  Tilapia      ....      403 

melanotheron,  Sarotherodon     .          .       389,  390 
Melitaea         ....  -47° 

meneliki,  Barbus     ....          .27-28 


496 


INDEX 


mento,  Barbus 
Merluccius 

merluccius,  Merluccius     . 
micra,  Woloszynskia 
microbarbis,  Barbus 
microcephala,  Tilapia 
microcephalus,  Brienomyrus 
microcephalus,  Mormyrus 
microcotyleum,  Lepidapedon 
Micromesistius 
microplax,  Amphiodia 
microplax,  Amphispina   . 
micropogon,  Chiloglanis  . 


154,  180-181 

154,  180-181 

.   194 

3,  87-89,  139,  145 

•  4°5 

•  337 
•   336,  338 

.   166 

154,  180-181 

461,  481 

461,  481 

33i,  358,  359,  360, 
361-365,  367,  413 

•  338 

72 
.  213 


microps,  Stomatorhinus  . 

Micropterus   .... 

Microscolex    .... 

microstoma,  Barbus  intermedius       .          .        48 

microterolepis,  Barbus    .  3,  44,  89-gi,  129-130, 

139,  M5 
.  36 
.  213 

3,  5,  91-94,  114,  130-131, 
133,  140,  145 
.  469 
.  480 
•  423 
331,  358-367 
426 
.   116 
154,  180-181 


miochilus,  Barbus  . 
minutus,  Rhododrilus 
mirabilis,  Barbus 


mirabilis,  Ophiothela 
mirabilis,  Spatagobrissus 
Mithrodia 
Mochokidae   . 
modica,  Decametra 
moeruensis,  Barbus 
Molva   . 


Monachocrinus 
monkei,  Auchenoglanis    . 
Monostoma    . 
montisarthuri,  Megascolex 


•  423 

•  358 

•  157 
204,  214,  319, 

320,  321-323,  324 
montisarthuri,  Perionychella  203,  206,  221, 

225,  230,  241,  244,  246-247,  254,  257 
montisarthuri,  Vesiculodrilus  203,  206,  221, 

225,  230,  241,  244,  246-247,  254,  257 
montiskosciuskoi,  Graliophilus  .  .  260 

moorii,  Barilius  .....  25 
morhua,  Gadus  .  .  .  .  .166 
Mormyridae  .  .  .  .  331,  334-338 
Mormyrus  .  331,  334-335,  336,  338,  413,  PI.  i 
moroea,  Diporochaeta  .  .  .  .257 
moroea,  Perichaeta  .  .  205,  223,  257 

moroea,  Perionychella  .  203,  206,  257-258 

mortoni,  Cryptodrilus  .  205,  219,  221,  247 
mortoni,  Perionychella  .  203,  206,  221,  223, 
225,  230,  244,  247-250,  254,  255,  257,  286, 

Pis.  99-102 

mortoni,  Plutellus 247 

mortoni,  Vesiculodrilus  .  203,  206,  221,  223, 

225,  230,  244,  247-250,  254,  257,  286, 
Pis.  99-102 

mortoni,  Woodwardiella ....  247 
multicolor,  Haplochromis  .  .  .  389 
multicolor,  Pseudocrenilabrus  .  .  .  389 
multifasciatus,  Sarotherodon  galilaeus  .  408 


multiprostatus,  Telmatodrilus . 
mungoensis,  Barbus      .         331, 

mungoensis,  Labeo 
Muraenesox   .          .          . 
myriaster,  Conger  . 
Myxotheca     .... 


205 

342,  343,  344- 
345-347,  413 

•  34i 

•  157 

•  157 

•  195 


nairobi,  Barbus  ....  • 94-9 5 
Nannaethiops  .  .  .  .  .411 
natalensis,  Amphiura  grandisquama  447-448 
natalensis,  Astropecten  granulatus  423,  431-433, 

PI.  i 

natalensis,  Barbus  .  .  .  .  .129 
nedgia,  Barbus  .  .  .9,  48,  50,  54-55 
nedgia,  Barbus  affinis  .  .  .  .48 
nedgia,  Labeobarbus  .  .  .  .  9,  48 
nemoralis,  Diporochaeta.  .  .  .  223 
Neosteganoderma  .  .  151,  154,  169-173 

neumayeri,  Barbus  ....       95 

neuvillii,  Barbus  .....  49 
Nicolla .  ....  151,  161-162 
nigra,  Tilapia  ....  72,  78 

nigrofasciatus,  Pelmatochromis  .  393,  394 
nigromarginatus,  Procatopus  .  .  380,  384 
niloticus,  Lates  .  .  .  .  .26 
niloticus  waterloti,  Chiloglanis  .  361,  363 

Notoscolex  204,  207,  263,  280,  303,  307,  310,  317 
nototaenia,  Procatopus  .  331,  376,  377,  379, 
380,  381,  382,  383,  384,  385-386 
novembris,  Gymnophiura  .  475,  476,  483 
novemradiata,  Anseropoda  .  .  .423 
nuda,  Ophiothela  .....  469 
nudum,  Ophiopsammium  .  .  469,  483 
nunensis,  Labeo  chariensis  .  .  .341 
nyongana,  Tilapia  .....  403 


obesus,  Barbus        .      10-11, 
obliquae,  Perionychella 


obliquae,  Vesiculodrilus 


16,  19,  70,  127-128 
203,  206,  221,  225, 
249,  250-251,  257 

2O3,  206,   221,   225, 
249,  250-251,   257 

obscura,  Perichaeta          .          .          .          .223 

obscurum,  Aphyosemion  .          .          .      375 

obscurus,  Channa   .          .          .          .          .411 

obscurus,  Chromidotilapia        .          .          .413 
obtusispina,  Perissasterias         .          .       440,  483 
occidentalis,  Barbus         ....      347 

occidentalis,  Oligometra  serripinna   .       424-426 
ochracea,  Melitaea  .....      470 

ochroleuca,  Amphichilus.          .          .          .461 

ochroleuca,  Amphioplus  .          .          .          .461 

Odinia  .......      482 

oeseri,  Aphyosemion        .         331,  332,  370,  371, 

373-376,  411,  412,  413 

oeseri,  Panchax       ....       373,  374 

officeri,  Cryptodrilus    204,  205,  208,  269,  280-282 


INDEX 


497 


officer!,  Notoscolex 

officer!,  Trinephrus 

Ogden,  C.  G. . 

ogowensis,  Tilapia  .          . 

Oligochaeta    . 

Oligometra     . 

olivacea,  Amphilimna 

Onchocerca    . 

Onos     .... 

Opecoelidae   . 

Ophiacantha . 

Ophiacanthidae 

Ophiactidae  . 

Ophiactis 

Ophiarachnella 

Ophidiaster    . 

Ophidiasteridae 

Ophiocirce 

Ophiocoma     . 

Ophiocomidae          .          . 

Ophioderma  . 

Ophiogymna . 

Ophioleucidae 

Ophiomastix  . 

Ophiomisidium 

Ophiomitrella 

Ophiomusium 

Ophiomyxa    . 

Ophionephthys 

Ophionereis    .          .          . 

Ophiopallas    . 

Ophiophragmus 

Ophiopsammium     . 

Ophiopsila 

Ophiopteron  . 

Ophioteresis  . 

Ophiothela     . 

Ophiothrix     . 

Ophiotrichidae 

Ophiura 

Ophiuridae     . 

Ophiuroidea  . 

opisthogaster,  Hickmaniella       204,  214,  270, 

300,  301-302,  324, 

Pis.  64-66 

oreas,  Barbus  ....  48,  57 

Oreasteridae  ......     435 

Oreoscolex      .   204,  205,  210,  212,  214,  280,  282, 

298,  302-318,  324, 
Pis.  67-90 

ornata,  Henricia     .....      439 
Otodistomum  ....       154,  167 

oviformis,  Gromia  .          .          .          .  195 

oxycephalus,  Barbus        .          .          .          .116 
oxyrhynchus,  Barbus  3,  36,  47,  50-51,  72,  78-80, 
82,  87,  94-IOO,  103,  127-129,  131,  140,  145 


pacei,  Anthosticte  . 
pacei,  Tethyaster    . 


280  Pachylometra          .....     429 

.     280  pagenstecheri,  Barbus    3,  101-IO3,  139-140,  145 

185-199,  Pis.  1-7  paludinosus,  Barbus         ....      129 

404,  405  Panchax         ....        373,  374,  375 

201-328,  102  Pis.  paradoxa,  Ophiopallas     .          .         423,  477-478 

423,  424-426  paradoxa  altera,  Ophiopallas    .          .       477,  478 

442,  444  Paranepanthia         .....     437 

332  parenzani,  Barbus  macronema  .  48,  52 

162  parva,  Ophiactis     ....       464,  465 

159-162  patagonicus  chondriscus,  Ceramaster          .     435 

442  patagonicus  euryplax,  Ceramaster     .       435,  483 

441-444  Patiriella        .....       438,  483 

464-465  paucispina,  Ophiopsila     ....     474 

464-465  paucisquamata,  Barbus  altianalis      .         76,  103 

480  paucisquamatus,  Barbus          3,  4,  n,  40-41,  76, 

.     436  103-io6,  140,  145 

435-436  pectinatus,  Amphioplus  .          .         456-459,  481 

423,  476-477  pectinatus,  Telmatodrilus          .          .          .      205 

472,  481  peculiaris,  Oreoscolex    .         204,  210,  303,  305, 

470-475  306,  313-315,  324,  Pis.  81-82 

423  pedderensis,  Perionychella    203,  206,  221,  225, 

•     423  244,  251,  257 

476-478  pedderensis,  Vesiculodrilus  203,  206,  221,  225, 

.     472  244,  251,  257 

476  pellegrini,  Geophagus       .          .          .  391 

441,  482  pellucida,  Trinema  lineare        .          .          .189 

476,  482  Pelmatochromis      .  388,  391,  393,  394,  402 

482  penardi,  Trinema    .          .          .          .          .189 

462-464,  483  pentaphylla,  Ophiothrix  fragilis         .          .     468 

.          .          .     480  pentaphyllum,  Ophiothrix  fragilis     .          .     468 

423,  477-478  percivali,  Barbus    .....       95 

452,  453,  454  Perichaeta     .  205,  217,  218,  220,  221,  222, 

469,  483  223,  228,  234,  257,  258,  259,  324 

470-475,  483  perince,  Barbus 353 

469  Perionychella    203,  204,  206,  209,  211,  212,  214, 

469,  483  216-259,  260,  263,  265,  266,  286,  Pis.  91-102 

469-470,  483  Perionychini  ....         203,  216-299 

465-469,  483  Perionyx        .          .  207,  217,  218,  221,  245 

465-470  Perissasterias          .          .          .         439-440,  483 

475-476,  483  perplexicans,  Barbus        .          .          -94,  96-97 

475-476  perplexicans,  Capoeta      ....       94 

440-479  perrieri,  Monachocrinus  ....     423 

petersi,  Gnathonemus  ....  338 
phalacrus,  Diporochaeta .  .  .  .221 
phalacrus,  Perionychella.  .  .  .216 
phalacrus,  Perionyx  .  .  .  .221 

Pheretima 217,  265 

pheretima,  Diporochaeta  .          .          .222 

phosphoreus,  Microscolex          .          .          .213 
Phoxaster      ......     434 

phragmorus,  Astropecten          .          .       433,  483 
phragmorus,  Astropecten  acanthifer.          .      433 
phragmorus,  Astropecten  polyacanthus  433-434, 

Pis.  1-2 
Phycis  ....  154,  162,  180-181 

pica,  Ophiocoma     .          .          .          .          .481 

pietschmanni,  Auchenoglanis   .          .       356,  357 
pietschmanni,  Barbus      .          .          .          .10-12 

423  Pinguidrilus         .    203,  204,  209,  212,  263-265, 

423  294,  324 


INDEX 


154.  J57 

1-148,  329-420,  5  Pis. 
.'         .       466,  467 


piscatorius,  Lophius 
Pisces    . 
Placophiothrix 
plagiostomus,  Barbus       .  .        48 

plana,  Ophiactis      .  .       464-465 

platyacanthus,  Amphioplus      .          .          .461 
platyrhinus,  Barbus        3,  44,  106-iog,  126-127, 

129,  139,  M5 
platystomus,  Barbus        .          .          .          .        48 

platystomus,  Barbus  platystomus     .          .        48 
platystomus  daga,  Barbus        .  .        48 

platystomus  dekkensis,  Barbus          .          .        48 
platystomus  platystomus,  Barbus     .          .        48 
platystomus  prognathus,  Barbus       .          .        48 
platystomus  vatovae,  Barbus  .          .  48,  65 

Plecostomus  ....  .      389 

Pleionogaster  ...  .      300 

plenum,  Benthotrema      .          .          .          .178 

pleuromelas,  Sarotherodon        .          .          .      409 
plumosus,  Procatopus      .  .       379,  380 

Plutellus         .    204,  207,  218,  222,  241,  247,  253, 
255,  257,  260,  263,  265,  266,  268 
pojeri,  Barbus         .....        36 

polyacantha,  Perissasterias       .         439-44O,  483 
polyacanthus,  Astropecten        .          .       433,  483 
polyacanthus,    Astropecten    polyacanthus    433- 

434,  PI.  2 

polyacanthus  phragmorus,  Astropecten  433-434, 

Pis.  1-2 

polyacanthus   polyacanthus,    Astropecten    433- 

434-  PL  2 

polyarthra,  Porphyrocrinus      .          .          .      482 
polylepis,  Barbus    .          .          .          .          .129 

Polymixia      .          .          .          .          .          .172 

polymixiae,  Neosteganoderma .          .       151,  172 

polymixiae,  Proctophantastes  .          .       151,  172 

polymorphus,  Amphioplus        .          .       463,  464 

polynephricus,  Cryptodrilus     204,  205,  267,  268, 

269,  270,  271,  273,  275, 

280,  282-283,  284,  286 

polynephricus,  Cryptodrilus  polynephricus    204, 

208,  281,  283,  284-288,  289, 

291,  292,  323,  Pis.  20-31 

polynephricus,  Trinephrus        .          .          .284 
polynephricus    ad    urethrae,     Cryptodrilus 
polynephricus         204,  208,  281,  286,  291-293, 

323,  Pis.  32-43 

polynephricus  polynephricus,  Cryptodrilus    204, 

208,  281,  283,  284-288,  289, 

291,  292,  323,  Pis.  20-31 

polynephricus  polynephricus  ad  urethrae, 

Cryptodrilus        .  204,  208,  281,  286,  291-293, 

323,  Pis.  32-43 

polynephricus  urethrae,  Cryptodrilus    204,  208, 

280,  281,  286,  287,  288-291,  292,  323, 

Pis.  44-59 

polyodon,  Chiloglanis       .          .         365,  366,  367 
Polypterus     .          .          .          .          .          .26 

Porphyrocrinus        .....      482 

porrecta,  Ophionereis       ....      480 


poutassou,  Micromesistius         .          154,  180-181 
pritchardi,  Megascolex     .          .          .       217,  220 
Procatopus    .     331,  332,  333,  376-384,  385-387, 

411,  412,  413 
procatopus,  Barbus          ....        49 

Proctophantastes    .          .  151,  167,  169-170,  172 
prognathus,  Barbus  platystomus       .          .        48 
Prosorhynchus         .  .  151,  154,  157-159 

proteus,  Acanthophiothrix        .         46S-4&6,  467 
proteus,  Ophiothrix          .  .         466-466,  467 

proteus,  Placophiothrix  .          .       •  .          .      466 
Protozoa        .          .          .          .  185-199 

Pseudocrenilabrus  .          .          .          .          .389 

Pseudocryptodi ilus  .          204,  205,  212,  214, 

296-299,  324,  Pis.  61-63 

pseudoexigua,  Patiriella  ....      438 

Psilaster         ......      434 

pulchellum,  Ophiomisidium      .          .          .      476 
pulchellum,  Ophiomusium        .          .          .476 
pumilus,  Phoxaster  ....      434 

Puntius  ......      349 

purpureus,  Vesiculodrilus          .          .          .      222 


quadrispinus,  Lophaster . 
queenslandica,  Rhododrilus 
quinquemaculata,  Ophiothrix 

rachiaea,  Lepodora 
rachion,  Distomum 
rachion,  Lepidapedon 
radcliffii,  Barbus     . 
radcliffii,  Barbus  altianalis 


.      438 

216 
•~ .       •      468 

162 

.      162 

151,  154,  162-I&4 

9-1 1,  23,  144 

9-1 1,  13,  15,  17, 


19-20,  22-27,  127,  129 

ramosa,  Stereomyxa  .  .  .  .194 
rarispina,  Marthasterias  glacialis  .  .  439 
rathkei,  Epiplatys  sexfasciatus  .  367,  369 

regani,  Pelmatochromis  ....  402 
reticulata,  Henricia  .  .  .  438-439 
reticulatus,  Echinaster  .  .  438,  439,  483 
rhinoceros,  Barbus.  ....  83 
rhipidiophorus,  Argulus  ....  35 
Rhizopodea  .  .  .  .  -  .187 
Rhododrilus  .  203,  204,  209,  212-2  16,  263,  297, 

324,  PI.  60 

richardi,  Diporochaeta  ....  222 
richardsoni,  Benthotrema  .  .  .178 
richea,  Diporochaeta  ....  258 
richea,  Perichaeta  .  .  .  2  05,  223,  258 
richea,  Perionychella  .  203,  206,  258-259 

roseipinnis,  Procatopus  .  .  .  .380 
rotunda,  Euglypha  .  187,  192,  i95-J97 

ruandae,  Varicorhinus      .          .        •  ';  87,  89 

ruasae,  Barbus  3,  7-9,  27,  73,  109-1  n,  128, 


rubra,  Megascolex  .          . 
rudolfianus,  Barbus  bynni 
rueppelli,  Barbus    .. 
rufoviride,  Lecithochirium 
rupestris,  Coryphaenoides 


.326 
.27-28 
48,  55 
178-179 
.169 


INDEX 


499 


rupestris,  Macrurus  . 

ruspolii,  Barbus      .          . 
rusticus,  Cryptodrilus 
ruwenzorii,  Varicorhinus 


169 

27-28,  35 

267,  269 

9,  "4 


saccarius,  Cryptodrilus    . 
sakaniae,  Barbus  chilotes 
salmoides,  Micropterus    . 
sanagaensis,  Tilapia 
santaisabellae,  Aphyosemion 
santa-isabellae,  Aphyosemion 
Sarotherodon 


•  303 

•  36 
72 

•  4°9 
373-  374.  376 

•  373 
331.  332,  388,  389,  390,  391,  404, 

405,  408-410,  411,  412,  413 

scheeli,  Aphyosemion  .  .  .  373,  376 
schoutdeni,  Chromidotilapia  .  .  .  397 
sclateri,  Antedon  .....  429 
sclateri,  Glyptometra  .  428,  429-430,  482 

sclateri,  Pachylometra  ....  429 
scolecoidea,  Diporochaeta  .  .  .259 
scolecoidea,  Perichaeta  .  205,  217,  223,  259 

scolecoidea,  Perionychella  203,  204,  206,  259 
scolecoides,  Perionychella  .  .  .  222 
Scomber  .....  154,  178 
scombrus,  Scomber  .  .  .  154,  178 

scutata,  Amphilepis  ....  464 
sedecimalis,  Diporochaeta  .  .  .223 
seguensis,  Barbus  .....  347 
seminuda,  Ophiopsila  .  .  476-472,  474 
semperi,  Ophiopsammium  .  .  .  469 
sentus,  Echinaster  .....  439 
sepositus,  Echinaster  ....  439 
serripinna,  Oligometra  .  .  .  423,  426 
serripinna  macrobrachius,  Oligometra  .  426 
serripinna  occidentalis,  Oligometra  .  424-426 
sexfasciatus,  Epiplatys  .331,  867-370,  411,  413 
sexfasciatus  rathkei,  Epiplatys  .  367,  369 

sexfasciatus  togolensis,  Epiplatys  .  .  369 
sexthecatus,  Oreoscolex  204,  210,  282,  303, 
305,  315-317,  324,  Pis.  83-86 


sigillatus,  Diporochaeta 
sigillatus,  Perionyx 
Silurus  . 

silvestris,  Hylopanchax 
similis,  Amphipholis 
similis,  Procatopus 


221 
221 

•      367 
.      382 

450,  451,  461 
33i.  332,  377.  378,  379, 


Simsia  . 

simsoni,  Amphiura 

simsoni,  Cryptodrilus 


simsoni,  Megascolides 

simulans,  Cryptodrilus     . 

Simulium 

sladeni,  Dipsacaster 

sladeni  capensis,  Dipsacaster 

Solasteridae   . 


somereni,  Barbus 
Sparus  . 


380,  381,  382,  383,  384-385,  386 

218 

448-449 

204,  208,  269,  270, 
273,  277,  293-296,  323 
205 

•  3°3 

•  332 

•  434 
.      434 

.      438 


Spatagobrissus 
spenceri,  Diporochaeta    . 
squamifera,  Ophiopsila    . 
stappersii,  Barbus  .        3, 
stappersii,  Varicorhinus  . 
steeli,  Megascolex   . 
Steganoderma 
Stelleroidea    . 
Stereomyxa   . 
Steringophoru  s 
Steringotrema 
Stomatorhinus 
strata,  Amphipholis 
striolata,  Ophiacantha     . 
sublineatus,  Barbus 
superba,  Hacelia     . 
superba  capensis,  Hacelia 
surkis,  Barbus    16,  47,  49, 
susanae,  Barbus 
svenssoni,  Barbus  . 
sylvaticus,  Megascolex     . 
Synodontis     . 


.      480 

•  223 

•  472 
116-1 19,  130,  139,  145 

•  36-37,  123 

326 

151,  154,  167-170,  172 
431-479 

•  194 

151,  154,  i74-J78 
151,  154,  173-174 

•  338 
450-452 

.     442 
352-355 

•  436 

•  435,  483 
51-52,  55,69,  127-128 

4.  M3 

•  356 
326 

26,  358 


Tamaria 
tanensis,  Barbus 
tanensis,  Lanceabarbus   . 
tanganicae,  Varicorhinus 
tanjilensis,  Cryptodrilus  . 
tanjilensis,  Perichaeta 
tapirus,  Mormyrus . 
Tasmaniaedrilus 
tasmaniaensis,  Tasmaniaedrilus 
tasmanianus,  Megascolides 
tasmanianus,  Notoscolex 
tasmanianus,  Pinguidrilus 


tasmanianus,  Plutellus 
tasmanica,  Perichaeta 
tasmanicus,  Megascolex 


•  436 
50,  72,  94-96,  100 

94 
126 

222 
22O 

331,  334-335,  413,  PI.  i 

•  205 

•  205 
263 

204,  207,  263 
203,  268-265, 

294,  324 

.  263 

205,  324 

204,  205,  214,  319,  320, 


3.  9,  92-94.  HI-US, 

130-131,  133,  140,  145 

.   408 


324-326 
Telmatodrilus          .....      205 

telopea,  Diporochaeta      ....      223 

tenuibrachia,  Comanthus  wahlbergi  .          .481 
tenuis,  Cryptodrilus          ....      269 

terrareginae,  Perionyx     .          .          .          .221 

terricola,  Trinema  lineare          .          .          .189 
tesselatus,  Cryptodrilus   .          .         205,  219,  266 
tesselatus,  Woodwardiella         .          .          .266 
tessellatus,  Plutellus         ....      266 

tessellatus,  Woodwardiella        .         203,  214,  266 
Testacea         .....        185-199 

Tethyaster     ......      423 

tetrastigma,  Barbus         ....      355 

Thalassometridae    ....        427-429 

tholloni,  Tilapia  .  .  .  389,  404,  405 
thysi,  Barbus  .  331,  350-356,  410,  411,  413 
Tilapia.  72,  78,  134,  331,  332,  388,  389,  391, 
402-408,  409,  411,  412,  413,  PI.  4 
timida,  Ophiopsila .....  475 


500 


INDEX 


tisdalli,  Megascolides  ....  222 
togolensis,  Epiplatys  sexfasciatus  .  .  369 
Tor  .  -134 

trachypterus,  Barbus  .  3,  40-41,  106,  116, 

119-123,140,  145 

Trachyrhynchus  .  154,  160-161,  164-166, 

167-168,  180-181 

trachyrincus,  Trachyrhynchus.  154,  160-161, 
164-166,  167-168,  180-181 

Trematoda 149-183 

Trewavas,  E.  329-420,  5  Pis. 

tricirratus,  Onos     .          .          .          .          .162 

trifolium,  Amphiophiura  .          .       423,  476 

triglochis,  Ophiothrix       .          .         468,  469,  483 
trimeni,  Ophiura     .          .          .         475-476,  483 
Trinema         .          .          .  185-199,  Pis.  1-7 

Trinephrus     ....         269,  280,  284 

tripapillatus,  Groliophilus      .       203,  214,  260, 

261-262,  324,  PI.  i 

trispilos,  Barbus     .          .  332,  350,  352-355 

trispinosus,  Ceramaster   .          .          .          .423 

Tritogenia      .          .          .          .          .          .301 

tropidolepis,  Barbus        3,  44,  108-109,  123-126, 

127,  129,  139,  145 
Tropiometra  ....         423,  426-427 

Tropiometridae       ....       426-427 

tunnackensis,  Perionychella  .  203,  206,  221, 

225,  230,  251,  252,  253,  257 
tunnackensis,  Vesiculodrilus  .  203,  206,  221, 

225,  230,  251,  252,  253,  257 
Tylochromis  .  .  .  .  .  .  391 


udekemianus,  Limnodrilus 

uncinatus,  Plutellus 

undulatum,  Lithodesium 

Unioplus 

unitaeniatus,  Nannaethiops 


•  205 
222 

.  194 
455-456 
411 


urethrae,  Cryptodrilus  polynephricus   204,  208, 

280,  281,  286,  287, 
288-291,  292,  323,  Pis.  44-59 
urethrae,  Cryptodrilus  polynephricus  poly- 
nephricus ad        .    204,  208,  281,  286,  291-293, 

323,  Pis.  32-43 

urundensis,  Barbus  .          .          .  in 

urundensis,  Barbus  alt ianalis   .          .       in,  113 


variegatus,  Lumbriculus ....  205 
vatovae,  Barbus  platystomus  .  .  48,  65 

veliporum,  Otodistomum  .          .       154,  167 

verrucosum,  Euryale  ....  440 
verrucosus,  Astrocladus  ....  440 
verrucosus,  Gorgonocephalus  .  .  .  440 
versluysii,  Barbus  .....  344 
Vesiculodrilus  203,  204,  206,  212,  216,  217,  218, 
219,  221-223,  225,  230,  231,  232,  233-253, 
254,  256,  257,  258,  263,  286,  Pis.  93-102 
vexillifer,  Aphyosemion  .  .  .  374,  375 
vexillifer,  Bathybiaster  ....  434 
vexillifer,  Panchax  .  .  373,  374,  375 

victoriae,  Barbus  .  .  .  .  .119 
victoriae,  Cryptodrilus  .  .  .  .222 
vivipara  capensis,  Ophiomyxa.  .  .  482 
volpinii,  Barbus  .  .  .  .  49,  52 

voltae,  Barbus  .....  356 
volvens,  Megascolides  .  .  .  .222 
vulgaris,  Gaidropsarus  .  .  .  154,  159 


wahlbergi,  Comanthus  ....  480 
wahlbergi,  Ophioderma  ....  423 
wahlbergi  tenuibrachia,  Comanthus  .  .481 
walhallae,  Perichaeta  .  .  .  .223 
walkeri,  Clarias  .  .  331,  356,  411,  413 
waterloti,  Chiloglanis  niloticus  .  361,  363 

weldboroughi,  Perionychella        203,  206,  220, 
223,  230-232,  256,  260 

wellingtonensis,  Cryptodrilus  .  .  205,  317 
wellingtonensis,  Notoscolex  .  .  -317 
wellingtonensis,  Oreoscolex  204,  210,  303,  305, 
316,  317-318,  324,  Pis.  87-90 
willsiensis,  Cryptodrilus  .  .  .  .222 
Woloszynskia  .  .  .  .  .194 
woodi,  Graliophilus  ....  260 
Woodwardia .  .  .  .  217,  219,  265 
Woodwardiella  .  203,  204,  209,  212,  214, 
217,  219,  247,  263,  265-266 


Xenodermichthys  . 


yunnanensis,  Barbus 


154.  174-176 


valida,  Anamphiura         .          .         423,  478-479  zaphiri,  Barbus 

Varicorhinus .       4,  9,  36-37,  49,  87,  89,  94,  114,  zillii,  Tilapia  . 

119-120,  123,  124,  126,  135  Zoogonidae    . 

varicus,  Derogenes.          .          .         154,  180,  181  zuaicus,  Barbus 


49 

403,  404,  405 

167-173 

49 


A  REVISION  OF  THE  LARGE 
BARBUS  (PISCES,  CYPRINIDAE) 
EAST  AND  CENTRAL  AFRICA 


K.  E.  BANISTER 


BULLETIN  OF 

THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 
ZOOLOGY  Vol.  26  No.  i 

LONDON:  1973 


A  REVISION  OF  THE  LARGE  BARBUsL* 
(PISCES,  CYPRINIDAE)  OF  EAST  AND  CENTRAL 

AFRICA  ' 

STUDIES  ON  AFRICAN  CYPRINIDAE 

PART  II 


JC 
KEITH  EDWARD  BANISTER/^ 


Pp.  1-148;  105  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
ZOOLOGY  Vol.  26  No.  i 

LONDON:  1973 


THE      BULLETIN      OF     THE      BRITISH      MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they 
become  ready.  Volumes  will  contain  about  three  or 
four  hundred  pages,  and  will  not  necessarily  be 
completed  within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  26  No.  i  of  the  Zoological  series. 
The  abbreviated  titles  of  periodicals  cited  follow  those 
of  the  World  List  of  Scientific  Periodicals. 


World  List  abbreviation : 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


Trustees  of  the  British  Museum  (Natural  History),  1973 


TRUSTEES   OF 
THE   BRITISH   MUSEUM   (NATURAL  HISTORY) 

Issued  i  November,  1973  Price  £6-95 


A  REVISION  OF  THE  LARGE  BARBUS 
(PISCES,  CYPRINIDAE)  OF  EAST  AND  CENTRAL 

AFRICA 

STUDIES  ON  AFRICAN  CYPRINIDAE 

PART  II 

By  KEITH  EDWARD  BANISTER 

CONTENTS 


SYNOPSIS  .......... 

INTRODUCTION  ......... 

Notes  on  counts  and  measurements  ..... 

Barbus  acuticeps         ......... 

Barbus  alluaudi  ......... 

Barbus  altianalis         ......... 

Barbus  bynni     .......... 

Barbus  caudovittatus  ......... 

Barbus  ethiopicus        ......... 

Barbus  gananensis      ......... 

Barbus  intermedius     ......... 

Barbus  longifilis          ......... 

Barbus  macrolepis       ......... 

Barbus  mariae  ......... 

Barbus  microbarbis     ......... 

Barbus  microterolepis  ........ 

Barbus  mirabilis         ......... 

Barbus  oxyrhynchus    ......... 

Barbus  pagenstecheri  ......... 

Barbus  paucisquamatus        ........ 

Barbus  platyrhinus     ......... 

Barbus  ruasae   .......... 

Barbus  somereni          ......... 

Barbus  stappersii        ......... 

Barbus  trachypterus    ......... 

Barbus  tropidolepis     ......... 

DISCUSSION       .......... 

ACKNOWLEDGEMENTS          ........ 

REFERENCES     .......... 

APPENDIX  i     An  artificial  key  to  the  species  described  in  this  paper 
APPENDIX  2     Gazetteer      ........ 

APPENDIX  3     A  replacement  name  for  Barbus  gregorii  Norman 
APPENDIX  4     Study  material       ....... 

INDEX 


4 
4 
5 
6 

9 
9 

27 
36 
41 
44 
47 
76 

79 
83 
87 
89 
9i 

94 
101 
103 
1 06 
109 
in 
116 
119 
123 
127 
135 
135 
139 
141 

143 

143 
145 


4  K.   E.    BANISTER 

SYNOPSIS 

,  il  the  previously  described  species  and  subspecies  of  the  large  Barbus  (the  species  usually 
described  as  possessing  parallel  striae  on  their  scales)  from  east  and  central  Africa  are  examined. 
The  area  under  consideration  is  bounded  in  the  north  by  the  mouth  of  the  Nile  and  in  the 
south  by  the  Zambezi  system.  The  western  boundary  is  the  upper  reaches  of  the  Congo  system 
adjacent  to  the  rift  valley  and  the  eastern  boundary  is  the  coast. 

It  is  shown  that  many  nominal  species  can  be  synonymized  when  a  sufficiently  large  series 
of  specimens  is  available  to  show  that  the  characters  formerly  used  to  distinguish  them  form 
a  continuous  series  within  a  polytypic  species.  Consideration  is  given  to  the  taxonomic  value 
of  these  characters  and  their  variability  is  demonstrated.  Particularly  variable  are  the  lips, 
body  depth  and  head  length.  This  is  a  reflection  both  of  eco-phenotypic  factors  and  of 
allometry. 

The  quaternary  palaeogeography  of  this  part  of  Africa  is  considered  in  an  attempt  to 
elucidate  the  present  distribution  of  certain  species. 

Two  former  subspecies  are  raised  to  specific  rank  (Barbus  longifilis  and  Barbus  paucisquamatus) 
and  one  new  subspecies  is  described  (Barbus  intermedius  australis).  Two  replacement  names 
are  included  ;  Barbus  lapsus  for  Barbus  babaulti  Pellegrin  1935  and  Barbus  susanae  for  Barbus 
gregorii  Norman  1923. 

INTRODUCTION 

THIS  revision  was  initiated  by  an  unsuccessful  attempt  to  identify  satisfactorily  the 
large  Barbus  species  collected  by  the  Sandhurst  Army  College  Ethiopian  Expeditions 
of  1964  and  1966.  The  use  of  the  keys  in  Boulenger's  Catalogue  of  African  Fresh- 
water Fishes  (1911-1916)  showed  that  some  species  were  so  imprecisely  denned  that 
an  individual  specimen  could  be  assigned  to  any  one  of  several  different  species. 
Many  of  the  species  described  by  early  workers  on  African  Barbus  species  were 
based  on  one  or  a  few  poorly  preserved  individuals,  and  it  is  only  now,  when 
sufficient  material  is  available,  that  an  attempt  can  be  made  to  redefine  some 
species  allowing  for  growth  and  natural  variation. 

Worthington  (i932a)  demonstrated  that  lip  shape,  formerly  considered  a  signifi- 
cant specific  character  in  the  large  Barbus  species,  was  extremely  variable.  This 
aspect  was  greatly  extended  by  Groenewald  (1958)  who  was  able  to  demonstrate 
the  great  range  in  Barbus  species  lip  form  engendered  by  the  environment  in  certain 
species.  In  the  course  of  this  work  he  cast  doubts  upon  the  validity  of  the  inclusion 
of  certain  species  in  the  closely  related  genus  Varicorhinus. 

Jubb  (1961,  1963,  1965,  I967a,  1968),  Crass  (1960)  and  Jackson  (1961)  have 
continued  this  study  and  have  partially  confirmed  Worthington's  (i932a)  views  on 
the  variability  of  the  large  Barbus  species.  Barbus  marequensis  A.  Smith  has  been 
shown  by  the  South  African  workers'  efforts  to  be  a  widespread  and  variable  species. 

An  analogous  situation  exists  in  eastern  and  central  Africa.  Many  large  Barbus 
species  have  been  described,  but  as  more  specimens  were  collected  the  boundaries 
between  the  species  became  obscured  as  new  data  rendered  the  old  specific  defi- 
nitions untenable.  New  descriptions  of  species  must,  therefore,  incorporate  the 
range  of  intraspecific  variation  if  the  definition  of  the  species  is  to  have  any  validity 
and  reflect  the  status  of  the  populations  in  the  field. 

Several  authors,  e.g.  Worthington  (i932a),  Pellegrin  (1935)  and  Bini  (1940),  have 
described  Barbus  subspecies  and  varieties.  The  subspecies  considered  in  this  paper 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  5 

conform  to  the  definition  of  Mayr  (1949)  although  it  has  not  always  been  possible 
to  conform  to  the  '75  per  cent  distinguishable'  convention  suggested  by  Mayr. 
Geographical  isolation,  on  its  own,  without  any  supporting  morphological  or  colour 
differences  is  not  considered  sufficient  justification  for  the  establishment  of  sub- 
species. I  would  like  to  have  used  Mayr's  definition  of  superspecies.  I  am  unable 
to  do  so  as  I  cannot  prove  that  the  species  I  consider  to  form  such  taxa  would 
constitute  a  monophyletic  assemblage  and  hence  I  use  the  terms  supra-specific 
complexes  or  groups.  This  does  not  assume  monophyly  although  hopefully  one 
day  it  may  be  proved. 

There  are  still  many  problems  that  I  have  been  unable  to  consider  in  this  work  ; 
ecological  information  is  only  rarely  available  (and  the  effect  of  the  environment 
could  well  be  an  important  factor  influencing  the  phenotypic  variation)  and  colour 
patterns  are  based  almost  entirely  on  preserved  specimens. 

This  revision  remains,  therefore,  essentially  that  of  a  museum  worker,  but  it  is 
very  possible  that  further  information  from  live  specimens  may  lead  to  the  estab- 
lishment of  subspecific  taxa  especially  in  the  widespread  species. 

There  are  some  geographical  areas  that  have  presented  many  difficulties,  par- 
ticularly where  watersheds  are  close  together.  By  unfortunate  quirks  of  geography 
or  international  politics  these  areas  are  inadequately  sampled  and  some  of  my 
subsequent  conclusions  about  the  status  of  the  specimens  must  be  regarded  as 
tentative.  These  problems  are  mentioned  in  the  text.  A  lot  of  time  was  spent  in 
trying  to  pin-point  the  localities  where  specimens  had  been  collected.  The  localities 
were  often  given  the  name  of  the  local  village,  but  these  villages  were  frequently 
only  temporary  settlements  of  nomadic  tribes.  Many  town  and  village  names  were 
changed  when  countries  were  occupied  or  invaded  and  collections  made  at  the  same 
site  under  different  regimes  bear  different  locality  names.  Where  such  localities 
have  been  traced  they  are  incorporated  in  the  gazetteer  (Appendix  2)  and  as  much 
information  as  I  can  give,  in  physiographical  details  or  in  grid  references,  is  included. 

The  limits  of  distribution  of  the  species  described  in  this  paper  are  from  the  Nile 
in  the  north  down  to,  but  not  including,  the  Zambezi  system.  The  eastern  limit 
is  the  eastern  coast  of  Africa  and  the  western  limit  is  the  western  rift  valley.  Some 
species  whose  range  extends  to  the  west  of  the  rift  valley,  into  the  Upper  Congo  are 
included,  e.g.  the  Upper  Congo  species  Barbus  mirabilis  is  included  because  of  its 
relationship  with  the  species  that  live  in  the  area  covered. 

The  arrangement  of  species  in  the  descriptive  part  of  the  text  is  alphabetical. 

Notes  on  counts  and  measurements 

The  standard  length  (S.L.)  was  taken  in  the  usual  manner  (see  Banister  1972). 
The  lateral  line  count  (LL)  was  taken  from  the  first  pore-bearing  scale  behind  the 
head  to  the  scale  lying  lateral  to  the  end  of  the  hypurals.  The  body  depth  (D)  is 
the  maximum  body  depth,  usually  to  be  found  just  in  front  of  the  dorsal  fin.  The 
anterior  limit  for  the  head  length  (H)  and  snout  length  (Snt)  was  the  premaxillary 
symphysis,  with  the  premaxillae  retracted.  In  'rubber-lipped'  fishes  this  measure- 
ment was  only  taken  with  difficulty.  The  posterior  limit  for  the  snout  is  the 


6  K.    E.    BANISTER 

anterior  margin  of  the  orbit.  The  term  mouth  width  (MW)  is  a  shorthand  notation 
for  the  width  of  the  lower  jaw  at  its  widest  point.  The  pectoral  fin  length  (Pet.) 
is  the  total  length  of  the  fin,  measured  in  a  straight  line  from  the  base  of  the  first 
ray  to  the  distal  extremity  of  the  fin.  The  measurement  is  taken  in  this  manner 
because  of  the  ease  of  so  doing  with  dial  calipers,  which  were  used  on  all  fish  except 
the  smallest  when  dividers  were  used.  The  caudal  peduncle  length  (CP1)  is  the 
horizontal  distance  from  the  posterior  angle  of  the  base  of  the  last  anal  fin  ray  to  the 
end  of  the  hypurals,  and  the  caudal  peduncle  depth  (CPd)  is  the  least  depth  of  that 
part.  The  interorbital  width  (10)  was  measured  as  the  least  distance  apart  of  the 
bony  edges  of  the  interorbital  space.  The  eye  diameter  (I)  had  to  be  taken  as  the 
horizontal  diameter  of  the  orbit  ignoring  the  skin  around  the  eye.  This  was  because 
in  many  of  the  more  ancient  specimens  the  skin  had  markedly  shrivelled  away  from 
the  eye  and  the  measurements  would  not  otherwise  have  been  comparable  with  those 
of  recently  preserved  fish.  The  dorsal  spine  (DSp),  strictly  the  last  unbranched  ray 
in  the  dorsal  fin,  was  measured  from  its  base  to  the  proximal  articulation  (if  any 
were  present).  The  reason  for  this  was  to  overcome  the  inaccuracy  caused  by 
damage  to  the  flexible  tips  and,  although  an  arbitrary  point,  experience  showed  it 
to  be  moderately  consistent.  Severely  damaged  spines  were  not  measured.  With 
any  measurements  which  were  repeatable  on  both  sides  of  the  fish  (e.g.  anterior 
barbel,  Ab,  or  posterior  barbel,  Pb),  the  larger  was  taken  except  in  a  few  cases  when 
gross  deformity  was  obvious. 

The  majority  of  measurements  were  taken  to  the  nearest  millimetre.  The 
exceptions  were  some  measurements  made  on  small  fishes  ;  these  were  taken  to  the 
nearest  half  millimetre.  The  limits  of  accuracy  do  not  justify  calculations  taken  to 
more  than  one  place  of  decimals.  The  mean  is  symbolised  by  x,  the  standard 
deviation  by  s.d.,  the  standard  error  by  s.e.  and  the  mode  by  m.  The  height  of  a 
pharyngeal  tooth  is  taken  as  the  parameter  at  90  degrees  to  the  plane  of  the  pharyn- 
geal  bone,  the  length  of  the  pharyngeal  tooth  is  the  parameter  of  a  transverse 
section  of  the  tooth  in  the  plane  of  the  anterior  edentulous  process  and  the  width  of 
the  tooth  is  the  parameter  at  90  degrees  to  the  length.  The  code  letters  for  the 
museums  from  whose  collections  the  specimens  came  are  listed  at  the  start  of 
Appendix  4. 

All  the  drawings  of  the  pharyngeal  bones  show  i)  the  bone  directly  from  above 
with  the  bone  lying  flat  on  its  edentulous  surface  and  2)  an  occlusal  view  of  the 
inner  row  of  teeth  with  the  bone  at  90  degrees  to  the  position  in  i),  i.e.  drawn  from 
above  with  the  bone  mounted  vertically  on  its  posterior  edentulous  process.  For 
convenience  these  are  referred  to  in  the  text  and  figure  captions  as,  respectively,  the 
dorsal  and  lateral  views. 

Barbus  acuticeps  Matthes  1959 

Barbus  acuticeps  Matthes,  1959,  Folia  scient.  Afr.  cent.  5  (3)  :  62  ;   Matthes,  1962,  Annls  Mus.  r. 
Afr.  cent.  Ser.  8vo  111  (2)  :  81,  pi.  i,  fig.  b. 

HOLOTYPE.  A  fish  of  210  mm  S.L.  from  the  Nyawarongo  river,  Lake  Rugwero, 
Rwanda.  Specimen  no.  M.A.C.T.  130313. 


A   REVISION   OF   THE   LARGE   AFRICAN   BARBUS 


FIG.  i.     Barbus  acuticeps  from  Matthes  1962. 


DESCRIPTION.  The  description  is  based  on  nine  specimens,  the  holotype,  four 
paratypes  (130310-12,  130314)  of  S.L.  43  mm,  121  mm,  92  mm  and  202  mm 
respectively,  and  four  other  specimens  M.A.C.T.  12910  (S.L.  131  mm),  13045-6 
(119  and  116  mm  S.L.)  and  172421  (242  mm  S.L.). 

My  standard  length  measurements  are  consistently  shorter  than  those  of  Matthes 
(1962).  I  attribute  this  to  different  concepts  of  standard  length. 


L 

D 

9 

H 

9 

I 

9 

10 

9 

MW 

8 

Pet 

8 

CP1 

9 

CPd 

9 

Snt 

8 

Ab 

8 

Pb 

8 

27-8 
28-7 

6-3 
7-1 

5-8 
21*7 
17-6 
n-4 

8-3 
4-8 
5-6 


s.d. 

4'i 
3'5 
i'3 
1-6 

i'3 

2-O 

1-7 

i'5 

1-2 
2-1 


s.e. 


0-7 
0-6 

o-5 
0-4 
0-8 
0-8 


range 
43    -242  mm 

23-6-33-7 
24-8-35-6 

4-3-  9-3 
4-7-10-4 

4-5-  7'9 
19-5-25-8 
15-2-20-6 

9-3-I3-7 
6-4-10-4 

2-2-    6-9 

2-9-  8-6 


All  measurements  are  expressed  as  percentages  of  the  standard  length. 

The  dorsal  profile  of  the  head  is  concave  and  a  pronounced  nuchal  hump  is  present. 
The  snout  is  pointed  and  the  mouth  slopes  upwards.  Otherwise  the  body  and  head 
shape  resembles  that  of  Barbus  altianalis  (sensu  lato)  and  Barbus  ruasae. 

Squamation.  It  was  not  possible  to  obtain  scale  counts  for  the  two  smallest 
specimens.  The  scales  have  parallel  striae  and  there  are  27  (1.3),  28  (f.i),  29  (f.2) 
or  30  (f.i)  scales  in  the  lateral  line.  Between  the  dorsal  mid-line  and  the  lateral 
line  there  are  4-5  (f.5)  or  5-5  (f.2)  scale  rows.  In  only  two  specimens  was  it  possible 
to  count  the  number  of  scale  rows  between  the  lateral  line  and  the  ventral  mid-line, 


8  K.    E.    BANISTER 

in  both  there  are  5-5  rows.  Between  the  lateral  line  and  the  base  of  the  pelvic  fin 
there  are  2  (f.2),  2-5  (f.2)  or  3  (f.i)  scale  rows.  There  are  12  scales  encircling  the 
caudal  peduncle. 

Dorsal  fin.  There  are  8  (f .5)  or  9  (£.3)  branched  rays.  The  fourth  unbranched 
ray  is  ossified  into  a  smooth,  straight  spine  (x  =  22-3  ;  s.d.  =  2-9  ;  s.e.  =  i-o  ; 
range  17-4-267).  There  is  no  sheath  of  scales  at  the  base  of  the  dorsal  fin.  The 
dorsal  fin  origin  is  slightly  behind  that  of  the  pelvic  fins.  The  anal  fin  has  three 
simple  rays  and  five  branched  rays. 

Gill  rakers.  In  five  specimens  examined,  three  had  eight  gill  rakers  on  the  lower 
limb  of  the  first  gill  arch,  the  other  two  had  ten  and  eleven. 

Pharyngeal  bones  and  teeth.  I  have  not  been  able  to  study  the  pharyngeal  bones 
but  they  were  described  by  Matthes  (1962)  as  '  Falciformes,  non  soudes,  attaches 
1'un  a  1'autre  par  une  symphyse  ligamenteuse.  Us  sont  peu  elargis  et  les  branches 
montantes  sont  comprimees  et  assez  greles.  Dents  pharyngiennes  obliquement 
tronquees  et  legerement  excavees,  avec  une  pointe  anterieur  recourbee  vers  1'arriere, 
en  3  rangees  au  nombre  de  2,  3,  5-5,  3,  2,  les  2me  et  3me  dents  de  la  rangee  interne 
un  peu  plus  fortes. ' 

Coloration.  Described  by  Matthes  (1962)  as  completely  silvery  with  pearly 
reflections  when  alive.  The  operculum  is  yellowish  with  metallic  glints,  the  eye  is 
yellowish,  the  pupil  ringed  with  gold.  The  fins  are  clear,  yellowish  ;  the  dorsal 
and  caudal  are  greyish,  yellowish  and  salmon-pink. 

DISTRIBUTION.  Four  of  the  specimens  came  from  the  Nyawarongo  river  (to  the 
north  of  Lake  Rugwero),  the  fifth  specimen  in  the  type  series  came  from  the  falls  of 
Rusumu  on  the  Kagera  (Rwanda).  The  other  specimens  came  from  Lake  Ihema, 
Kagera  ;  the  Nyabugogo  river  (an  affluent  of  the  Lusine  river,  Burundi)  and  from 
Matale  (?  Burundi). 

DIAGNOSIS  AND  AFFINITIES.  >Barbus  acuticeps  is  dubiously  distinct  from  Barbus 
altianalis.  The  principal  differences  between  Barbus  acuticeps  and  Barbus  altianalis 
are  i)  the  concave  dorsal  profile  of  the  head  and  the  nuchal  hump  of  the  former 
species,  2)  the  antero-dorsal  gape  of  the  mouth  of  the  former  species  which  contrasts 
with  the,  usually,  subterminal  mouth  in  Barbus  altianalis,  3)  fewer  gill  rakers  in 
Barbus  acuticeps  (8-n)  compared  with  10-14  (most  frequently  11-13)  m  Barbus 
altianalis,  4)  slightly  fewer  scales  in  the  lateral  line  series  in  Barbus  acuticeps  (27-30) 
compared  with  28-36  (most  frequently  30-34)  in  Barbus  altianalis,  5)  a  longer 
dorsal  fin  spine  in  Barbus  acuticeps  (x  =  22-3,  range  17-4-26-7)  compared  with 
x  =  16-9,  range  9-0-30-4,  in  Barbus  altianalis. 

It  must  be  remembered  that  Barbus  acuticeps  is  known  from  very  few  specimens 
and  that  more  are  necessary  to  confirm  whether  or  not  its  continued  separation  from 
Barbus  altianalis  is  justified. 

Barbus  ruasae  (p.  109)  also  from  Rwanda  to  some  extent  resembles  Barbus 
acuticeps  in  general  appearance  but  is  readily  separable  on  the  lower  number  of 
scales  in  the  lateral  line  series  [25  (f-9),  26  (f.i)].  The  pharyngeal  teeth  of  Barbus 
acuticeps  are  less  molariform  than  those  of  Barbus  ruasae,  but  this  could  be  the 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  g 

result  of  diet.  Barbus  ruasae  does  not  have  a  mouth  opening  antero-dorsally. 
Barbus  acuticeps  is  considered  here  to  belong  to  the  Barbus  intermedium  group  (see 
p.  128). 

Barbus  alluaudi  Pellegrin,  1909 

Barbus  alluaudi  Pellegrin,  1909,  Bull.  Soc.  zool.  Fr.  34  :  155  ;    Pellegrin,  1910,  Mem.  Soc.  zool. 
Fr.  22  :  287,  pi.  14,  fig.  2  ;    Banister,  1972,  Bull.  BY.  Mus.  nat.  Hist.  (Zool.)  24  (5)  :  261-290. 


1cm 


FIG.  2.     Barbus  alluaudi  Holotype. 


This  specific  name  has,  I  believe,  been  given  to  hybrids  between  Barbus  somereni 
and  Varicorhinus  ruwenzorii  [Banister  (1972)]. 


Barbus  altianalis  Boulenger,  1900 

Barbus  altianalis  Blgr.,  1900,  Ann.  Mag.  nat.  Hist.  (7)  6  :  479  ;    Blgr.,  1901,  Trans,  zool.  Soc. 

London,  16  :  144,  pi.  13,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  36  ;   Norman,  1925,  Occ.  Pap. 

Boston  Soc.  nat.  Hist.  5  :  189. 
Barbus  altianalis  altianalis  :   Worthington,  1932,  /.  Linn.  Soc.  (Zool.)  38  (258)  :  124-127  (Lake 

Kivu  and  Ruzizi  river  specimens)  ;   Poll,  1953,  Result,  scient.  Explor.  hybrobiol.  Lac  Tanganika 

(3)  5A  :  88. 

Barbus  altianalis  radcliffii  :   Worthington,  1932,  ibid.  :  124-127  (Lake  Victoria  specimens). 
Barbus  altianalis  eduardianus  :   Worthington,  1932,  ibid.  :  124-127  (Lakes  Edward  and  George 

specimens) . 
Barbus  altianalis  var.  labiosa  Pellegrin,  1935,  Revue  Zool.  Bot.  afr.  28  (3)  :  376-385  (only  the 

holotype,  from  Lake  Kivu). 
Barbus   (Labeobarbus)   nedgia  :    Hilgendorf,   1888,   Sber.  Ges.  naturf.  Freunde  Berl.  :  88   (from 

Lake  Victoria,  misidentification,  not  Barbus  nedgia  Riippell ;  fide  Blgr.,  1911). 
Barbus  eduardianus  Blgr.,  1901,  Ann.  Mag.  nat.  Hist.  (7)  8  :  12  ;    Blgr.,  1907,  Fish  Nile  :  230, 

pi.  43,  fig.  2. 
Barbus  fergussonii  Blgr.,   1901,  Ann.  Mag.  nat.  Hist.  (7)  8:12;    Blgr.,  1907,  Fish  Nile  :  230, 

pi.  43,  fig.  i. 

Barbus  radcliffii  Blgr.,  1903,  Ann.  Mag.  nat.  Hist.  (7)  12  :  218  ;    Blgr.,  1907,  Fish  Nile  :  212, 
pi.  37,  fig.  i. 


io  K.   E.    BANISTER 

Barbus  marequensis  :   Pellegrin,  1904,  Mem.  Soc.  zool.  Fr.  17  :  17  (misidentification,  not  Barbus 

marequensis  Smith,  specimen  from  Kavirondo,  Lake  Victoria,  fide  Blgr.,  1911). 
Barbus  lobogenys  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  435  ;    Blgr.,  1907,  Fish  Nile  :  210, 

pi.  36,  fig.  i. 

Barbus  bayoni  Blgr.,  1911,  Annali  Mus.  civ.  Stor.  nat.  Giacomo  Doria  (3)  5  :  77. 
Barbus  kivuensis  Pappenheim,  1914,  Wiss.  Ergebn.  dt.  ZentAfr.  Exped.  Zool.  (3)  5  :  237. 
Barbus  gregorii  :   Blgr.,  1916,  Cat.  Afr.  Fish  4  :  230  (only  the  specimens  from  the  Malawa  river). 
Barbus  pietschmanni  Lohberger,  1929,  Anz.  Akad.  Wiss.  Wien  66  :  92-94. 
Barbus  hollyi  Lohberger,  1929,  Anz.  Akad.  Wiss.  Wien  66  :  92-94. 
Barbus  obesus  Worthington,  1929,  Proc.  zool.  Soc.  Lond.  (3)  :  433. 
Barbus  kiogae  Worthington,  1929,  Proc.  zool.  Soc.  Lond.  (3)  :  434. 
Barbus  longirostris  Worthington,  1929,  Proc.  zool.  Soc.  Lond.  (3)  :  435. 

NOTES  ON  THE  SYNONYMY.  Barbus  altianalis  is  a  widespread  species  occupying 
three  isolated  areas.  The  fish  of  each  of  these  areas,  although  in  morphometric 
characters  scarcely  different  modally  from  one  another,  were  given  subspecific 
status  by  Worthington  (iQ32a). 

Barbus  altianalis  was  first  described  from  two  fish  from  Lake  Kivu  and  one  from 
the  Ruzizi  river.  All  are  large  specimens  ;  the  smallest  (from  the  Ruzizi)  has  a 
S.L.  of  306  mm  while  the  other  two  are  331  and  406  mm. 

Barbus  eduardianus  was  described  from  one  large  fish  from  Lake  Edward  and 
Barbus  fergussonii  from  two  small  fishes  from  the  same  lake.  The  latter  two 
specimens  have  unusually  shallow  bodies  when  compared  with  equal-sized  specimens 
even  from  the  same  waters. 

Barbus  radcliffii  was  based  on  a  skin  from  Lake  Victoria.  Barbus  lobogenys  was 
based  on  two  'rubber-lipped'  specimens  from  Lake  Victoria.  Barbus  bayoni  was 
described  from  a  very  large  fish  (S.L.  640  mm)  from  near  Jinja  on  Lake  Victoria. 
Barbus  pietschmanni  and  Barbus  hollyi,  both  from  Lake  Victoria,  were  based 
respectively  on  one  and  two  specimens. 

The  holotype  and  sole  example  of  Barbus  obesus  from  Lake  Kioga  (fig.  3)  is  a  fat, 
deep-bodied  fish  with  a  low  number  of  lateral  line  scales  (28).  The  scales  are 
slightly  lobed  and  the  specimen  gives  the  impression  of  a  rather  deformed  individual. 
The  two  syntypes  of  Barbus  longirostris  (fig.  4)  from  the  same  locality  are  large 
specimens  (388  and  410  mm  S.L.)  with  shallow  bodies  and  heads  longer  than  the 
mean  for  the  species.  They  are  linked  with  more  typical  Barbus  altianalis  by 
members  of  the  type  series  of  Barbus  kiogae.  No  morphometric  or  meristic  way 
can  be  found  for  separating  the  populations  from  Lake  Kioga  from  those  of  Lake 
Victoria.  Large  fish  from  Lake  Victoria  not  infrequently  approximate  to  the  facies 
of  Barbus  longirostris  and  Barbus  kiogae  (P.  H.  Greenwood  :  pers.  comm.).  The 
description  of  the  holotype  of  Barbus  kivuensis  differs  hardly  at  all  from  that  of 
Barbus  altianalis  from  the  same  lake. 

Norman  (1925)  commented  that  the  specimens  of  Barbus  altianalis  from  Lake 
Edward,  collected  by  Dr  John  Phillips,  were  identical  with  Barbus  radcliffii  from 
Lake  Victoria. 

Worthington  (iQ32a)  was  able  to  gather  together  sufficient  material  to  realize  that 
a  single  species  is  found  in  the  three  areas,  Barbus  altianalis  altianalis  from  Lake 
Kivu  and  the  Ruzizi  river  ;  Barbus  altianalis  radcliffii  from  Lake  Victoria  and  the 


A   REVISION   OF   THE   LARGE   AFRICAN   BARBUS 


ii 


FIG.  3.     Barbus  altianalis  (Holotype  of  Barbus  obesus  re-drawn  from  Worthington  1929). 


Victoria  Nile  ;  and  Barbus  altianalis  eduardianus  from  Lakes  Edward  and  George 
and  the  Kazinga  channel. 

Boulenger's  Barbus  raddiffii,  Barbus  lobogenys  and  Barbus  bayoni  were  synony- 
mized  by  Worthington  (i932a)  in  Barbus  altianalis  raddiffii,  and  Barbus  eduardianus 
and  Barbus  fergussonii  in  Barbus  altianalis  eduardianus.  An  important  conclusion 
of  Worthington's  work  was  the  realization  that  Barbus  species  can  exhibit  remark- 
able intraspecific  variation,  particularly  with  regard  to  the  development  of  the  lips 
and  he  suggested  that  further  extensive  synonymizing  might  be  necessary  when 
larger  samples  became  available.  However,  he  did  not  synonymize  the  species  he 
described  from  Lake  Kioga  because  with  the  material  at  his  disposal  they  remained 
apparently  distinct.  Scatter  diagrams  comparing  body  depth  and  head  length  of  the 
Lake  Victoria  and  Lake  Kioga  populations  are  shown  below  (figs.  6  and  7).  Pel- 
legrin  (1933,  1935)  described  several  varieties  of  Barbus  altianalis  from  Kivu  region  : 
only  one  of  these  is  considered  here  as  belonging  to  this  species  (for  the  others  see 
under  Barbus  caudovittatus,  Barbus  paucisquamatus  and  Barbus  longifilis] .  A  slight 
element  of  confusion  was  introduced  by  Pellegrin's  use  of  'Region  du  Kivu'  ;  this 
is  not  always  the  Lake  Kivu  basin  but  in  some  cases  refers  to  sites  in  the  Congo 
system. 

Greenwood  (1966)  synonymized  Barbus  pietschmanni  and  Barbus  holly i  with 
Barbus  altianalis  raddiffii. 

I  am  retaining  Worthington's  three  subspecies,  certainly  on  geographical  grounds 
but  also  because  of  the  consistent  colour  differences  between  the  Barbus  subspecies 
of  Lake  Victoria  and  those  of  Lake  Edward-George.  Dr  P.  H.  Greenwood  (pers. 
comm.)  has  noted  that  specimens  of  Barbus  altianalis  eduardianus  of  over  100  mm 
S.L.  from  Lake  Edward-George  have  pink  caudal  fins  whereas  those  of  Lake  Victoria 
never  do.  I  have  no  information  about  the  specimens  from  Lake  Kivu. 


12 


K.   E.    BANISTER 


FIG.  4.     Barbus  altianalis  (figured  specimen  of  Barbus  longirostris, 
re-drawn  from  Worthington  1929). 


FIG.  5.     Barbus  altianalis  altianalis  (Holotype  from  Boulenger  191  la). 


LECTOTYPE.  A  fish  of  331  mm  S.L.  from  Lake  Kivu  (one  of  the  three  in 
Boulenger's  type  series).  B.M.  (N.H.)  Reg.  No.  1906.9.6  :  13. 

DESCRIPTION.  The  description  is  based  on  a  total  of  213  specimens  ;  15  of  these 
are  from  Lake  Kivu  and  the  Ruzizi  river  (size  range  46-410  mm  S.L.),  130  are  from 
Lakes  Edward  and  George,  the  Kazinga  channel  and  Lake  George  feeder  streams 
(size  range  68-436  mm)  and  68  are  from  Lakes  Victoria  and  Kioga  (size  range 
99-640  mm).  The  combined  morphometric  data  for  all  three  populations  are  as 
follows,  and  include  all  the  holotypes  except  for  Barbus  pietschmanni  and  Barbus 
hollyi. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


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x  s.d.  s.e.  range 

L  46    -640  mm 

D  29-4  2-4  0-4  22-4-37-4 

H  26-1  1-6  o-i  22-5-31-4 

I  7-6  i-o  o-i  4-1-11-5 

IO  8-4  i-o  0-7  6-4-11-2 

MW  6-2  0-9  o-i  4-7-  9-5 

Pet  21-8  1-4  o-i  17-5-25-0 

CP1  16-9  1-4  o-i  13-1-19-8 

CPd  12-0  0-9  o-i  9-5-16-2 

Snt  8-3  0-8  o-i  5-8-10-9 

Ab  4-3  i-i  o-i  2-1-11-6 

Pb  5-4  1-3  o-i  2-3-12-4 

The  body  shape  is  extremely  variable,  especially  in  large  fish.  This  is  more 
noticeable  in  Lake  Kioga  than  elsewhere  and  a  graph  showing  the  variation  in  body 
depth  to  length  can  be  seen  in  fig.  7.  It  is  possible  that  the  deeper-bodied  specimens 
are  females  but  most  of  the  specimens  that  I  have  seen  had  been  gutted  (see  below). 

The  mouth  is  sub-terminal  and  usually  in  the  shape  of  a  narrow  horse-shoe. 
'Rubber-lipped'  forms  have  been  found  (e.g.  the  holotype  of  Barbus  lobogenys  -  see 
fig.  8).  The  snout  tends  to  overhang  the  mouth  in  fishes  of  medium  size. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays.  The  last  unbranched  ray 
is  enlarged  into  a  smooth,  straight  or  slightly  curved,  well-ossified  spine  with  articu- 
lations only  at  the  tip.  The  dorsal  fin  spine  is  negatively  allometric  ;  x  =  16-9, 
s.d.  =  3-2,  s.e.  =  0-2,  range  =  9-0-30-4  (percentage  of  the  S.L.)  for  the  whole 
sample  (fig.  9).  This  range  is  too  large  to  be  useful  as  a  diagnostic  character  but  it 
can  be  seen  from  fig.  9  that  there  are  three  convenient  size  ranges  of  fishes  which  can 
be  treated  separately  :  0-170  mm,  x  =  19-2,  s.d.  =  3-0,  s.e.  =  0-3,  range  =  13-7- 
30-4  ;  171-250  mm  S.L.,  x  =  16-8,  s.d.  =  1-7,  s.e.  =  0-2,  range  =  12-8-20-6  ; 
above  251  mm  S.L.,  x  =  13-4,  s.d.  =  2-3,  s.e.  =  0-3,  range  =  9-9-20-2.  It  is 
interesting  to  note  that  these  three  S.L.  ranges  are  the  same  for  Barbus  intermedius 
(see  p.  68). 

The  branched  rays  number  8  (1.30),  9  (f.i77)  or  10  (f.6).  The  relative  propor- 
tions of  fishes  with  8,  9  or  10  rays  does  not  differ  detectably  in  the  three  subspecies. 
The  dorsal  fin  origin  is  situated  in  front  (rare)  or  more  commonly  directly  above  or 
just  behind  the  vertical  from  the  origin  of  the  pelvic  fins. 

The  anal  fin  has  three  simple  rays  and  five  branched  rays  ;  the  unbranched  rays, 
although  thickened,  are  much  more  flexible  than  those  in  the  dorsal  fin. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  teeth  almost  always  number 
2.3.5.  ~  5-3-2-  The  one  exception  to  this  is  a  specimen  from  Lake  George,  S.L. 
208  mm  B.M.  (N.H.)  Reg.  No.  1971.8.11  :  33-39  which  is  abnormal  and  the  teeth 
number  1.1.2.3.5.  ~  5-3-2.1. 

The  pharyngeal  bones  may  vary  in  thickness  in  fishes  of  the  same  size.  The 
variability  in  the  length  and  thickness  of  the  bones  and  the  teeth  is  a  striking  feature 
of  this  species  and  Barbus  intermedius.  Both  species  have  remarkably  variable 
body  forms  and  there  appears  to  be  a  loose  correlation  between  body  depth  and  the 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  8.     Barbus  altianalis  radcliffii  (Type  of  Barbus  lobogenys  from  Boulenger  191  la). 


..••  v 

~  JV 

•    ••» 


40     8°     100     2°      40     60      80     200     20     40      60      80     300     20     40      60     80     400     20     4°      60      80     500     20     40     60      80    600     20      *°      6° 

FIG.  9.  Scatter  diagram  of  the  dorsal  spine  length  as  a  percentage  of  the  standard  length 
against  the  standard  length  for  Barbus  altianalis.  (=  Barbus  altianalis  altianalis, 
Barbus  altianalis  radcliffii,  Barbus  altianalis  eduardianus) . 


16 


K.    E.    BANISTER 


FIG.  10.     Dorsal  and  lateral  views  of  the  left  pharyngeal  bone  of  the  holotype  of 

Barbus  altianalis  altianalis. 


tip-to-tip  length  of  the  pharyngeal  bone.  The  very  deep-bodied  specimen  of  Barbus 
altianalis  (i.e.  Barbus  obesus)  and  the  deep-bodied  Barbus  intermedius  (i.e.  Barbus 
surkis]  both  have  pharyngeal  bones  that  are  very  much  longer  and  stouter  than 
those  of  less  deep-bodied  fishes  of  the  same  standard  length. 

A  series  of  ten  fishes  from  Lake  Victoria  and  ten  from  Lakes  Edward  and  George 
were  examined  to  see  what  relationship  was  present  between  the  length  or  depth  of 
the  body  and  the  size  of  the  pharyngeal  bone.  The  results  from  this  small  trial 
sample  must  be  used  with  considerable  restraint  but  using  the  coefficient  of  variation 
technique  there  are  indications  that  the  length  of  the  pharyngeal  bone  is  closely 
related  to  the  depth  of  the  body.  I  do  not  want  to  pursue  this  matter  any  further 
or  in  any  detail  but  simply  mention  the  possibility  that  this  correlation  is  worth 
further  study. 

The  pharyngeal  bone  of  a  long-headed  specimen  (Barbus  longirostris]  is  shown  in 
fig.  12. 

Diet  presumably  must  have  an  effect  upon  the  stoutness  of  the  pharyngeal  bones 
and  upon  the  shape  of  the  teeth.  The  stomach  of  the  deep-bodied  Barbus  inter- 
medius mentioned  above  (and  p.  69)  is  full  of  gastropod  mollusc  shells,  which,  if 
this  were  its  normal  diet,  could  explain  the  molariform  teeth  and  the  stout  bone  as 
having  been  developed  as  a  result  of  breaking  open  strong  shells.  Regrettably,  the 
stomach  contents  of  the  deep-bodied  Barbus  altianalis  were  unidentifiable  and  there 
were  no  traces  at  all  of  shells.  The  extent  to  which  diet  affects  the  pharyngeal  teeth 


A    REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  1 1 .     Dorsal  and  lateral  views  of  the  left  pharygeal  bone  of  a  specimen  of 
Barbus  altianalis  eduardianus. 


FIG.  12.     Dorsal  and  lateral  views  of  the  left  pharyngeal  bone  of  Barbus  altianalis 
radcliffii  (one  of  the  types  of  Barbus  longirostris  S.L.  410  mm). 


18 


K.   E.   BANISTER 


A-F.J. 


10mm 


FIG.  13.  A  series  of  pharyngeal  bones  from  specimens  of  Barbus  altianalis  eduardianus, 
to  show  the  variation  in  shape  with  the  growth  of  the  fish.  The  standard  lengths  (mm) 
of  the  specimens  are  :  A  =  90  ;  B  =  100  ;  C  =  101  ;  D  =  125  ;  E  =  141  ;  F  =  171  ; 
G  =  205  ;  H  =  227  ;  I  =  258  ;  ]  =  330. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  14.  A  series  of  pharyngeal  bones  from  specimens  of  Barbus  altianalis  radcliffii. 
Compare  with  the  series  opposite.  Standard  lengths  (mm)  of  the  specimens  are  :  A  =  84  : 
B  =  102  ;  C  =  118  ;  D  =  125  ;  E  =  145  ;  F  =  160  ;  G  =  197  ;  H  =  228  ;  I  =  255  ; 
J  =  343;  K  =  410;  L  =  388.  K  is  from  the  holotype  of  Barbus  longirostris  ;  L  is  from 
the  holotype  of  Barbus  obesus. 


20  K.   E.    BANISTER 

in  these  species  cannot  be  determined  at  the  moment,  but  further  considerations 
are  mentioned  on  p.  127. 

Consistent  differences  were  noted  in  the  pharyngeal  bones  and  teeth  of  specimens 
of  Barbus  altianalis  radcliffii  and  Barbus  altianalis  eduardianus  of  the  same  size 
(figs.  13  and  14).  The  pharyngeal  bones  of  Barbus  altianalis  radcliffii  are  consis- 
tently stouter  than  those  of  Barbus  altianalis  eduardianus  and  the  angle  of  the 
anterior  process  is  different. 

About  15  per  cent  of  the  pharyngeal  bones  examined  had  a  molariform  second 
tooth  in  the  inner  row.  This  phenomenon  is  as  common  in  Lake  Victoria  as  it  is  in 
Lakes  Edward  and  George  and  is  apparently  unrelated  to  size,  the  molariform  tooth 
being  found  in  a  specimen  of  only  121  mm  S.L. 

The  change  in  the  shape  of  the  pharyngeal  bones  and  teeth  with  the  increase  in 
the  size  of  the  fish  can  be  seen  in  figs.  13  and  14.  These  should  be  compared  with 
the  series  of  pharyngeal  bones  of  Barbus  bynni  (fig.  19)  -  a  species  with  a  less  variable 
body  form. 

Cyprinid  pharyngeal  bones  and  teeth  have  frequently  been  considered  good 
characters  for  distinguishing  species.  The  value  of  this  character  must  now  be 
reconsidered  in  the  light  of  the  observed  variability  of  these  characters  in  Barbus 
altianalis.  There  are  two  possible  interpretations  of  the  evidence  :  that  either  the 
degree  of  variation  in  the  pharyngeal  bones  and  teeth  has  not  been  fully  realized 
and  that  it  is  a  dubious  character  at  the  species  level ;  or  that  certain  species  only 
have  variable  pharyngeal  bones  and  teeth  and  that  for  these  species  the  very 
variation  is  a  good  character.  Insufficient  evidence  is  available  at  the  moment 
to  expand  the  arguments  for  either  of  the  two  interpretations. 

Squamation.  The  lateral  line  follows  a  gentle  curve.  There  are  from  28  to  36 
scales  in  the  lateral  line  ;  28  (£.3),  29  (f.6),  30  (f.2o),  31  (f.4i),  32  (£.57),  33  (£.41), 
34  (f.26),  35  (f.g),  36  (f.2)  (fig.  59).  The  number  of  scales  in  the  lateral  line  series 
could  not  be  counted  on  all  the  specimens.  There  are  6-5  (less  frequently  5-5, 
rarely  4-5)  scales  between  the  dorsal  mid-line  and  the  lateral  line  and  5-5  (less 
frequently  6-5,  rarely  4-5)  scales  between  the  lateral  line  and  the  ventral  mid-line. 
The  number  of  scale  rows  between  the  lateral  line  and  the  origin  of  the  pelvic  fin 
varies,  between  2  and  3-5.  The  number  of  scales  around  the  caudal  peduncle 
varies  from  12  to  14. 

Gill  rakers.  There  are  from  10  to  14  (most  frequently  11-13)  gill  rakers  on  the 
lower  limb  of  the  first  gill  arch  in  all  three  subspecies. 

Coloration.  Generally  the  body  colour  in  live  fish  is  from  a  silvery-grey  to  a 
bronzy-green,  the  scales  frequently  having  dark  bases,  the  operculum  slightly 
bronzy.  In  fresh  specimens  from  Lake  George  the  pectoral  fins  are  faintly  pink 
with  a  darker  posterior  half.  The  pelvics  are  grey  suffused  with  pink  ;  the  anal  fin 
membrane  is  dark  at  the  base.  The  caudal  fin  is  salmon-pink  with  a  dark  posterior 
edge  in  specimens  of  over  100  mm  S.L.  from  Lakes  Edward  and  George.  This  is  a 
diagnostic  character  separating  the  Lake  Victoria  and  the  Lakes  Edward  and 
George  subspecies. 


A   REVISION  OF  THE   LARGE   AFRICAN  BARBUS                               21 

Barbus  altianalis  altianalis  Boulenger 

A  general  description  of  the  species  is  given  on  p.  12.     For  comparative  purposes 

morphometric  data  of  the  15  specimens  (S.L.  46-410  mm)  from  Lake  Kivu  and  the 
Ruzizi  river  are  given  below. 

x  s.d.                  s.e.  range 

L  49    -410  mm 

D                     26-9  2-3                  0-6  22-6-30-3 

H                     25-6  1-4                  0-4  23-4-28-0 

I                        7-0  2-1                  0-5  4-5-10-2 

IO                     7-8  1-2                  0-3  6-6-10-3 

MW                  6-0  0-8                  0-2  4-6-  7-7 

Pet                  22-0  i-i                   0-3  20-2-23-8 

CP1                  15-6  1-3                  0-3  13-1-17-6 

CPd                 n-o  0-7                  0-2  10-1-11-4 

Snt                    8-5  0-9                  0-2  7-3-10-1 

Ab                     4-5  i-i                   0-3  2-4-  6-6 

Pb                     5-9  1-2                  0-3  4-6-  8-3 

DSp                16-9  2-3                  0-6  13-3-23-1 

Lateral  line  count  30  (1.5)  ;  31  (f.2)  ;   32  (f.3)  ;  33  (f.3)  ;  34  (f.2). 


Barbus  altianalis  eduardianus  Boulenger 

HOLOTYPE.  A  fish  of  371  mm  S.L.  B.M.  (N.H.)  Reg.  No.  1906.9.7  :  41  from 
Lake  Edward,  Uganda. 

DESCRIPTION.  The  description  is  based  on  130  specimens.  A  general  descrip- 
tion is  given  above.  The  morphometric  data  in  detail  are  as  follows. 

x  s.d.  s.e.  range 

L  68    -436  mm 

D  29-6  2-i  0-2  22-4-37-0 

H  25-8  1-6  o-i  22-5-31-4 

I  8-0  1-5  o-i  4-1-11-5 

IO  8-1  0-9  o-i  6-4-10-8 

MW  6-0  0-7  o-i  4-4-  8-3 

Pet  22-1  1-3  o-i  18-8-25-0 

CP1  17-1  1-2  o-i  14-4-19-7 

CPd  12-4  0-9  o-i  10-1-16-2 

Snt  8-1  0-7  o-i  5-8-10-1 

Ab  4-2  0-7  o-i  2-2-  6-4 

Pb  5-4  0-9  o-i  3-7-  7-3 

DSp  17-5  2-4  0-2  11-7-24-0 

Lateral  line  count  28  (f.i),  29  (f.5),  30  (f.g),  31  (f.2i),  32  (f.39),  33  (1.22), 
34  (f.i7)  35  (f.6),  36  (f.2).  The  lateral  line  scales  could  not  be  counted  on  all  the 
specimens  examined. 


K.   E.    BANISTER 


It  is  perhaps  unwise  to  compare  this  sample  with  the  much  smaller  sample  from 
Lake  Kivu  but  the  general  agreement  in  mean  values  indicates  that  the  two  popu- 
lations are  extremely  similar. 


Barbus  altianalis  radcliffii  Boulenger 

HOLOTYPE.  A  skin  of  530  mm  S.L.  B.M.  (N.H.)  Reg.  No.  1904.5.19  :  13  from 
Lake  Victoria. 

DESCRIPTION.  The  description  is  based  on  68  specimens  (S.L.  90-640  mm).  A 
general  description  is  given  above.  The  morphometric  data  in  detail  are  given 
below. 

x  s.d.  s.e.  range 

L  90    -640  mm 

D  29-6  2-7  0-3  24-0-37-4 

H  26-7  1-7  0-2  23-8-29-8 

I  7-0  i-i  o-i  4-1-  9-3 

IO  9-0  i-o  o-i  7-5-11-2 

MW  6-6  i-o  o-i  4-7-  9-5 

Pet  21-3  1-6  0-2  17-5-24-6 

CP1  16-8  1-7  0-2  13-2-19-8 

CPd  11-7  0-9  o-i  9'5-13'Q 

Snt  8-7  0-7  o-i  7'5-io-g 

Ab  4-5  1-6  0-2  2-1-11-6 

Pb  5-3  1-9  0-2  2-3-12-4 

DSp  15-9  4-2  0-5  9-0-30-4 

Lateral  line  count  28  (f.2),  29  (f.i),  30  (f.6),  31  (f.i8),  32  (f.i5),  33  (f.i6), 
34  (f-7)>  35  (f-3). 

The  mean  values  for  the  measurements  taken  differ  little  from  those  of  the  other 
two  populations.  The  absence  of  the  red/pink  caudal  fin  in  Barbus  altianalis 
radcliffii  has  been  mentioned  above. 

DISTRIBUTION.  Barbus  altianalis  is  found  in  Lake  Kivu  and  the  adjacent  part 
of  the  Ruzizi  river,  Lake  Victoria  including  the  Lake  Kioga  basin  and  Lakes  Edward 
and  George.  Although  Lake  Victoria  and  Lakes  Edward  and  George  are  still  con- 
nected, migration  along  the  connecting  waterways  is  not  easy  at  the  moment.  The 
presence  of  Barbus  altianalis  in  Lake  Kivu,  which  is  isolated  from  Lake  Victoria 
and  Lakes  Edward  and  George,  and  its  absence  from  Lake  Albert  are  facts  which 
must  be  explained.  A  certain  amount  is  known  about  the  geological  history  of  the 
lakes  and  I  have  attempted  to  assemble  and  correlate  recorded  events  from  each 
lake  and  to  try  to  explain  the  distribution  of  this  species. 

During  the  early  Pliocene,  the  region  now  occupied  by  Lake  Victoria  was  drained 
to  the  west  by  the  upper  tributaries  of  the  Congo  system  (Cooke  1958).  This 
drainage  system  persisted  well  into  the  Pleistocene  (Kendall  1969).  Stages  in  the 
formation  of  the  western  rift  valley  interrupted  this  flow  and  the  upwarp  of  the 


A   REVISION   OF   THE   LARGE   AFRICAN   BARBUS 


FIG.  15.     Barbus  altianalis  radcliffii  (the  type  of  Barbus  radcliffii, 
from  Boulenger  191  za). 


FIG.  16.     Map  of  the  distribution  of  the  three  subspecies  of  Barbus  altianalis  :  B.  altianalis 
altianalis  =  %,  B.  altianalis  eduardianus  =  A,  B.  altianalis  radcliffii  =  |. 


24  K.    E.  BANISTER 

eastern  shoulder  of  the  rift  caused  the  rivers  to  reverse  their  flow  and  pond-up  in 
the  shallow  depression  between  the  eastern  and  western  rifts.  This  led  to  the  for- 
mation of  the  present-day  Lake  Victoria  which  Temple  (1969)  dates  from  the  mid- 
Pleistocene.  Tectonic  movements  have  altered  the  lake  since  its  original  formation. 

Lacustrine  deposits  at  Nsongezi  in  the  Kagera  valley  (to  the  west  of  Lake  Vic- 
toria) are  about  35  m  above  the  present  lake  surface,  and  some  no  km  inland  of  its 
western  shore.  The  dates  of  the  Nsongezi  deposits  are  in  doubt.  Kendall  (1969) 
cites  authors  who  have  given  dates  ranging  from  60  ooo  years  B.P.  to  10  ooo  years 
B.P.  Doornkamp  and  Temple  (1966)  suggest  that  continued  tectonic  warping 
has  caused  the  eastward  drift  of  the  western  shore  of  the  lake.  Climatic  events 
have  probably  had  an  effect  as  well.  Kendall  (1969)  states  that  from  before 
14  500  years  B.P.  to  about  12  ooo  years  B.P.  Lake  Victoria  was  a  closed  basin  and 
the  water  had  a  high  concentration  of  salts.  From  about  12  ooo  years  B.P.  to 
about  10  ooo  years  B.P.  the  lake  had  an  exit,  which  could  have  been  to  the  south, 
but  its  locality  is  by  no  means  certain.  Around  10  ooo  B.P.  to  about  9500  B.P.  the 
basin  became  closed  again  and  then  drained  to  the  north  cutting  the  Nile  gorge. 
The  water  level  in  the  lake  rose  after  12  500  B.P.  There  followed  a  period  of  aridity 
around  10  ooo  B.P.,  the  water  level  fell  and  the  outlet  was  lost.  A  wetter  climate 
ensued,  the  lake  level  rose,  overflowed  to  the  north  and  established  the  Nile  outflow. 
It  is  difficult  to  correlate  these  events  because  tectonic  movements,  climatic  changes 
and  the  rate  of  discharge  from  the  lake  have  all  had  an  effect  upon  the  lake  size, 
level  and  position.  The  date  of  cessation  of  the  tectonic  movements  is  not  known 
with  certainty  but  there  are  three  horizontal  raised  beaches  3,  12  and  18  m  above 
the  lake  level  which  were  presumably  formed  after  earth  movements  had  ceased. 
The  most  recent  of  these  has  been  dated  at  3720  years  B.P.  (Stuvier  et  alii  1960). 
Kendall  (1969)  argues  that  they  all  must  be  post  12  500  B.P.  which  may  well  indi- 
cate that  tectonicity  had  stopped  by  that  time.  The  different  beach  levels  are 
possibly  related  to  stages  in  the  erosion  of  the  Nile  gorge. 

Much  less  is  known  about  Lake  Kivu  and  very  little  absolute  dating  has  been 
done.  Lake  Kivu  resulted  from  the  effects  of  eruptions  of  the  Mufumbiro  (or 
Bufumbiro)  volcanoes  (to  the  north  of  the  present  lake)  when  lava  flows  dammed 
the  rift  valley.  Prior  to  this  unknown  date  the  drainage  in  this  part  of  the  rift  is 
usually  thought  to  have  been  northwards,  but  there  may  have  been  a  proto-Lake 
Kivu.  Cahen  (1954)  describes  the  origin  of  Lake  Kivu  as  the  result  of  the  landscape 
changes  which  affected  a  network  of  rivers  flowing  S.W.  to  N.E.  which  curved 
northwards  and  ponded-up  to  form  the  lake.  This  direction  of  flow  is  possibly  not 
compatible  with  the  ideas  of  Cooke  (1958)  who  describes  the  drainage  as  roughly 
E.  to  W.  but  this  may  have  been  due  to  local  topographical  features.  The  lake  later 
rose  in  height  and  spilled  over  southwards  to  drain  into  Lake  Tanganyika  cutting 
the  Ruzizi  gorge.  Kendall  (1969)  noted  that  circa  12  450  B.P.  Lake  Kivu  was 
100  m  higher  than  its  present  level  This  date  was  established  by  radio-carbon 
dating  on  mollusc  shells  on  a  raised  beach.  Possibly  this  immediately  pre-dates 
the  formation  of  the  Ruzizi  river. 

The  critical  date  of  the  eruption  of  the  Mufumbiro  volcanoes  is  unknown.  Cahen 
(1954)  states  that  it  was  before  the  end  of  the  alleged  Kamasien  ;  Bishop  (pers. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  25 

comm.)  dates  the  eruptions  as  upper  Pleistocene  and  Fryer  (1968)  suggests  a  date 
of  about  100  ooo  years  B.P.  Lake  Kivu  has  been  connected  to  Lake  Tanganyika 
for  a  minimum  period  of  12  ooo  years  and  although  Barbus  altianalis  is  found  in  the 
Ruzizi  river  at  the  Lake  Kivu  end  it  has  not  been  recorded  from  Lake  Tanganyika 
(Poll  1953  ;  Marlier  1953).  Barilius  moorii  has  made  the  journey  in  the  other 
direction  (Poll  1950).  Why  only  Barilius  moorii  has  been  able  to  ascend  the  Ruzizi 
river  is  a  question  that  I  am  unable  to  answer.  It  may  be  that  the  ascent  of  the 
river  is  easier  for  some  species  than  the  descent  is  for  others.  There  are  also  rapids 
on  the  Ruzizi  river  which  must  limit  the  amount  of  faunal  interchange.  All  that 
one  can  say  at  the  moment  is  that  for  at  least  the  last  12  ooo  years  only  one  species 
of  fish  is  known  to  have  successfully  navigated  the  Ruzizi  river. 

A  lake  occupying  the  site  on  the  floor  of  the  western  rift  valley  now  occupied  by 
Lakes  Edward  and  George  began  to  form  at  the  time  of  the  uplift  of  the  rift  wall. 
This  corresponds  with  fig.  2B  in  Doornkamp  and  Temple  (1966),  and  Bishop  (1971) 
dates  it  as  late  Pliocene.  Rivers  drained  onto  the  floor  of  the  rift  and  a  lake 
resulted.  There  was  probably  more  than  one  lake,  for  the  extensive  Kaiso  beds 
(later  Pliocene)  are  lacustrine  as,  largely,  are  the  overlying  Semliki  deposits  (mid-  to 
later  Pleistocene) .  The  full  extent  of  the  '  Kaiso '  and  '  Semliki '  lakes  both  in  time 
and  area  is  not  completely  known.  The  Kaiso  beds  are  up  to  8000  feet  thick 
(Brown  1956)  and  represent  possibly  three  million  years  of  comparative  stability. 
Bishop  (1969)  noted  that  the  Kaiso  and  Semliki  deposits  are  separated  in  time  by 
renewed  movement  of  the  rift  valley  boundary  faults.  The  Semliki  beds  are  found 
north  of  Lake  Albert  indicating  a  very  large  area  for  the  '  Semliki'  lake.  The  upper- 
most Semliki  beds  are  later  Pleistocene  with  an  age  in  excess  of  10  ooo  years  B.P. 

At  the  end  of  the  Semliki  period  soil  formation  occurred  and  about  8000  years 
to  10  ooo  years  B.P.  the  Katwe  volcanoes  ceased  erupting  and  the  present  forms  of 
Lakes  Edward,  George  and  the  Kazinga  channel  became  defined.  The  Kazinga 
channel  runs  through  the  Katwe  lavas,  but  probably  dates  from  the  time  that  soil 
formation  occurred,  i.e.  the  post-Semliki  period.  The  form  of  the  channel  was 
little  altered  by  the  volcanic  activity  (Bishop  1969  :  45). 

The  Ishango  beds,  which  were  deposited  during  the  active  phase  of  the  Katwe 
volcanoes  (8000  years  to  10  ooo  years  B.P.,  Bishop,  op.  cit.},  have  yielded  fossil 
Barbus  species  remains  at  Ishango  on  the  Semliki  river  exit  from  Lake  Edward 
(Greenwood  1959) .  From  Site  IX  in  Makalian  strata,  fossils  comparable  with  Barbus 
bynni  and  Barbus  altianalis,  then  dated  Mesolithic  or  Holocene,  were  described  by 
Greenwood  (op.  cit.}.  Site  VI  at  Ishango,  in  the  upper  Semliki  beds  (Katanda  for- 
mation), dated  mid-Pleistocene  [but  older  than  the  Nsongezi  deposits  (Bishop  1969)], 
yielded  unidentifiable  Barbus  species  fragments,  the  oldest  known  from  this  area. 

Today  the  fish  fauna  of  Lake  Edward  is  different  from  that  of  Lake  Albert,  the 
former  lacking  species  present  in  the  latter.  The  Semliki  river  connects  the  two 
lakes  ;  probably  the  rapids  and  possibly  the  cooler  streams  flowing  down  from  the 
Ruwenzori  mountains  (Fryer  1968)  prevent  any  faunal  interchange.  The  fauna 
of  Lake  Edward  is  poor,  lacking  representatives  of  some  widespread  African  fish 
families,  e.g.  Polypteridae,  Characidae,  Mochokidae,  Centropomidae  and  Masta- 
cembelidae,  as  well  as  some  specifically  Nilotic  species  present  in  Lake  Albert,  e.g. 


26  K.    E.    BANISTER 

Lates  niloticus  and  Barbus  bynni.  This  situation  has  been  described  in  detail  by 
Greenwood  (1959). 

Some  of  the  species  now  absent  from  Lake  Edward  have  been  found  fossilized  in 
the  Ishango  beds.  Kendall  (1969)  states  that  the  major  fish  extinctions  probably 
occurred  within  the  last  18  ooo  years.  Polypterus,  Lates,  Synodontis  and  Barbus  cf. 
bynni  were  present  some  8000-10  ooo  years  ago.  Characins  disappeared  much 
earlier  in  the  fossil  record,  the  latest  known  fossil  Hydrocynus  occurring  in  the  Kaiso 
beds  (probably  lower  Pleistocene).  Certainly,  at  the  time  the  Ishango  beds  were 
deposited,  the  fish  fauna  of  Lake  Edward  was  closer  to  that  now  in  Lake  Albert 
than  it  is  today. 

The  Ishango  beds  lie  on,  and  are  also  covered  by,  volcanic  rocks  (Bishop  1969). 
So,  it  would  seem  very  possible  that  some  species  were  killed  by  the  outpourings  of 
the  Katwe  eruptions.  It  is  less  likely  that  a  drying  up  of  the  lake  caused  the 
extinctions  because  there  is  no  evidence  for  such  an  event  and  Kendall  (1969) 
suggests  that  in  the  Lake  Victoria  basin  the  period  from  9500  B.P.  to  6500  B.P.  was 
marked  by  high  water  levels  and  the  levels  fell  only  slightly  from  6500  B.P.  onwards. 

It  is  uncertain  whether  Lake  Edward  became  completely  denuded  of  fish  life  at 
the  end  of  the  Ishango  period.  If  it  did  then  the  lake  has  been  recolonized  ;  if  it 
did  not,  then  some  species  must  have  escaped,  or  must  have  remained  unaffected  by 
the  eruptions.  Dr  P.  H.  Greenwood  (pers.  comm.)  reports  having  seen  fish  behaving 
normally  within  a  hundred  metres  of  a  lava  stream  entering  Lake  Kivu,  this  suggests 
that  perhaps  an  increase  of  toxic  products  in  the  water  is  more  lethal  than  the  heat- 
ing effect  of  the  lava.  Possible  means  by  which  various  species  could  have  survived 
are  discussed  by  Greenwood  (1959).  They  include  migration  into  feeder  streams  and 
different  degrees  of  resistance  to  toxicity  and  low  oxygen  concentrations. 

If  there  was  an  invasion  from  Lake  Victoria  it  must  have  occurred  since  8000  B.P. 
(the  date  of  cessations  of  the  Katwe  volcanoes).  However,  since  that  time  there  is 
no  evidence  that  Lake  Victoria  has  been  subjected  to  any  great  earth  movements 
(see  above)  and  it  had  already 'gained  its  outlet  via  the  Nile  gorge.  This  would 
limit  any  rises  in  water  level  to  about  18  m  above  the  present  level.  The  watershed 
between  Lakes  Edward  and  George  and  Lake  Victoria  is  extremely  low.  Now  the 
Katonga  river  is  blocked  by  Papyrus  swamp  but  a  few  metres  rise  in  water  level 
might  possibly  enable  fishes  to  travel  from  Lake  Victoria  into  Lakes  George  and 
Edward.  The  fact  that  no,  e.g.  Haplochromis,  fossils  have  been  found  in  the 
Ishango  beds  does  not  necessarily  mean  that  they  have  invaded  the  lake  in  post- 
Ishango  times.  The  fossils  could  yet  await  discovery.  Present  studies  (Dr  P.  H. 
Greenwood,  pers.  comm.)  have  suggested  that  Lake  Edward  Haplochromis  species 
are  not  as  closely  related  to  particular  Lake  Victoria  Haplochromis  species  as  had 
been  thought,  which  possibly  militates  against  the  idea  of  an  invasion  from  Lake 
Victoria. 

It  seems  much  more  likely  that  the  fauna  of  Lakes  Edward  and  George  and  Lake 
Kivu  was  originally  widespread  in  the  upper  reaches  of  the  old  Congo  system  and 
became  isolated  in  the  present  lake  basins  as  a  result  of  tectonicity.  It  would  be 
very  difficult  to  explain  satisfactorily  how  Barbus  altianalis  could  have  migrated 
into  Lake  Kivu  from  either  Lake  Edward  or  Lake  Victoria.  At  the  time  of  the 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  27 

highest  known  level  reached  by  Lake  Kivu,  12  450  B.P.,  Lake  Victoria  was  not  at 
its  highest.  However,  it  must  be  noted  that  Lake  Victoria  possibly  had  an  outlet 
(see  above)  whose  direction  is  unknown.  The  general  topography  of  the  Lake  Kivu 
basin  makes  invasion  from  the  east  or  north  extremely  unlikely. 

If,  as  judged  by  their  breeding  habits  (Whitehead  1964),  Barbus  species  are 
fundamentally  fluviatile  fishes,  then  it  is  more  likely  that  Barbus  altianalis  originally 
lived  in  the  upper  reaches  of  the  Congo  system  in  the  area  now  occupied  by  Lakes 
Kivu,  Edward,  George  and  Victoria.  Since  the  suggested  modifications  in  its  range 
the  Barbus  altianalis  populations  have  undergone  no  significant  morphological 
changes  but  the  colour  difference  in  the  caudal  fin  is  noted  above. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  altianalis  has  three  recognized  subspecies 
which  are  meristically  and  morphometrically  very  similar.  The  pink  caudal  fin 
in  Lakes  Edward  and  George  subspecies  Barbus  altianalis  eduardinanus  separates 
this  subspecies  from  Barbus  altianalis  radcliffii  of  Lake  Victoria.  No  information 
is  available  on  the  live  colour  of  Barbus  altianalis  altianalis  from  Lake  Kivu. 

Barbus  altianalis  is,  in  gross  morphology,  similar  to  Barbus  intermedius  but  the 
two  species  are  separable  on  the  modal  lateral  line  count  (see  fig.  59),  26-34  (most 
frequently  28-32  ;  mode  29)  in  Barbus  intermedius  ;  28-35  (most  frequently  30-34, 
mode  32)  in  Barbus  altianalis.  The  last  unbranched  ray  in  the  dorsal  fin  is  slightly 
shorter  in  Barbus  altianalis  (9-0-30-4  per  cent  S.L.,  x  =  16-9)  than  in  Barbus  inter- 
medius (6-9-33-9,  x  =  20-1).  The  anterior  and  posterior  barbels  are  shorter  in 
Barbus  altianalis  (Ab  2-1-11-6  per  cent  S.L.,  x  =  4-3  ;  Pb  2-3-12-4,  x  =  5-4)  than 
in  Barbus  intermedius  (Ab  i-8-n-i,  x  =  5-8  ;  Pb  1-9-12-1,  x  =  6-9). 

For  a  comparison  of  Barbus  acuticeps  and  Barbus  altianalis  see  under  the  former 
species. 

Barbus  ruasae,  from  Rwanda,  is  somewhat  similar  in  appearance  to  Barbus 
altianalis  but  can  be  distinguished  easily  from  it  by  the  fewer  lateral  line  scales 
(25-26  against  28-35). 

Barbus  altianalis  is  considered  here  to  belong  to  the  Barbus  intermedius  group. 


Barbus  bynni  (Forsskal)  1775 

Cyprinus  bynni  Forsskal,  1775,  De scrip.  Anim.  :  71. 

Barbus  bynni  :    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  26,  fig.  8  (includes  a  full  bibliography  up  to  this 

date). 

Barbus  ruspolii  Vinciguerra,  1896,  Annali  Mus.  civ.  Stor.  natn.  Giacomo  Doria  (2)  17  :  29. 
Barbus  meneliki  Pellegrin,  1905,  Bull.  Mus.  Hist.  nat.  Paris  16  :  293 
Barbus  bynni  vudolfianus  Worthington,  1932,  /.  Linn.  Soc.  (Zool.)  38  :  132. 

NOTES  ON  THE  SYNONYMY.  The  type  specimen  of  Barbus  bynni  is  no  longer 
extant  (Klausewitz  &  Nielsen  1965)  and  a  neotype  has  been  selected  (see  below). 
The  holotype  of  Barbus  ruspolii  is  in  a  very  poor  state  and  has  not  been  seen  by  me, 
but  other  specimens  referred  by  Vinciguerra  to  Barbus  ruspolii  have  been  examined. 
The  holotype  and  unique  specimen  of  Barbus  meneliki  in  the  Paris  Museum  has  been 
studied. 


28 


K.   E.   BANISTER 


FIG.  17.     Barbus  bynni  (from  Boulenger  191  la). 


The  similarity  of  Barbus  bynni  to  Barbus  ruspolii  was  noted  by  Boulenger  (1907) 
and  by  Worthington  (i932a).  The  latter  author  was  also  of  the  opinion  that  Barbus 
meneliki  from  Lake  Rudolf  was  close  to  his  Barbus  bynni  rudolfianus.  Barbus 
meneliki  is  very  similar  in  many  respects  to  Barbus  bynni  rudolfianus  from  the  same 
lake.  The  greatest  difference  is  the  presence  of  29/30  lateral  line  scales  in  the  former 
against  34  or  35  in  the  latter.  Barbus  bynni  from  Lake  Albert  have  a  lateral  line 
range  of  28  to  35  scales.  Barbus  meneliki  is  based  on  the  holotype  and  unique 
specimen  so  either  it  is  an  extremely  rare  fish  in  the  lake  or  it  is  a  member  of  Lake 
Rudolf's  Barbus  bynni  population  with  fewer  scales  than  usual.  As  I  have  seen 
only  five  Barbus  bynni  specimens  from  Lake  Rudolf  the  latter  suggestion  seems  the 
more  likely  especially  when  the  information  is  compared  with  the  lateral  line  scale 
count  range  in  Lake  Albert  fish.  The  mouth  of  Barbus  meneliki  has  a  horny  covering 
on  the  lower  jaw,  although  this  has  not  been  seen  in  other  Lake  Rudolf  specimens  of 
Barbus  bynni,  it  is  not  an  unusual  phenomenon  in  Barbus  species  (Groenewald 
1958).  In  all  other  respects  (e.g.  dorsal  spine  length,  caudal  peduncle  shape  and 
scale  striations)  Barbus  meneliki  and  Barbus  ruspolii  both  closely  resemble  Barbus 
bynni  hence  I  consider  them  conspecific.  Boulenger  (1907)  remarked  that  little 
work  had  been  done  on  the  morphometric  variation  in  Barbus  bynni,  both  within 
one  population  and  between  isolated  populations.  What  little  I  have  been  able  to 
do  is  detailed  below. 

NEOTYPE.  Klausewitz  and  Nielsen  (1965)  listed  Cyprinus  bynni  as  being  'no 
longer  in  existence  from  Forsskal's  original  collection'.  I  have  also  tried  but 
without  success  to  trace  this  specimen  and  must  agree  with  Klausewitz  and  Nielsen 
that  the  specimen  is,  at  least,  lost.  Therefore  I  have  selected  as  a  neotype 
B.M.  (N.H.)  specimen  No.  1907.12.2  :  1230,  a  fish  of  152  mm  S.L.  collected  from 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


29 


the  Nile  at  Asswan.  Forsskal  did  not  state  the  size  of  his  specimen  nor  its  exact 
locality,  so  I  have  selected  this  fish  as  it  appears  to  be  a  modal  representative  of  the 
fishes  from  the  lower  Nile. 

DESCRIPTION.  The  description  is  based  on  a  total  of  63  fish  ;  36  from  the  lower 
and  White  Niles  ;  ten  from  Lake  Albert ;  five  from  the  Blue  Nile  ;  seven  from 
Lake  Abaya  and  five  from  Lake  Rudolf.  A  further  specimen  doubtfully  from  the 
Webi  Shebeli  will  be  treated  separately. 

The  body  is  shaped  like  an  elongate  rhomboid,  the  dorsal  profile  rises  sharply  and 
evenly  from  the  snout  to  the  occiput  and  then  more  steeply  to  the  origin  of  the  dorsal 
fin.  The  profile  of  the  belly  slopes  down  to  a  point  in  front  of  the  ventral  fins. 
The  body  is  compressed,  more  so  than  in  most  other  species  under  consideration. 
The  compressed  caudal  peduncle  is  short  and  deep  and  is  quite  characteristic  for 
Barbus  bynni. 

The  scales  bear  sinuous  longitudinal  striae. 

The  most  distinctive  features  are  the  massive  dorsal  spine  and  the  caudal 
peduncle.  At  least  one  specimen  (a  fish  from  the  lower  Nile)  shows  the  development 
of  'rubber  lips'. 

The  following  tables  show  the  range,  mean,  standard  deviation  and  standard 
error  of  the  measurements  taken  for  the  five  populations. 

Range  of  standard  lengths  of  the  populations  examined  (in  mm) 


No.  of  specimens 

Nile 

59-351 

36 

Lake  Albert 

136-349 

10 

Blue  Nile 

95-246 

5 

Lake  Abaya 

236-329 

7 

Lake  Rudolf 

196-233 

5 

Total 

59-351 

63 

Body  depth  (this  and  all  subsequent  data  are  expressed  as  percentages  of  the  S.L.) 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


n 

36 
10 

5 
7 
5 

63 


32-9 
34'6 
34'3 
35-o 
33-6 

33-6 


s.d. 

2-5 
2-7 
1-9 

2'5 

2-3 
2-5 


s.e. 

0-4 
0-8 
0-8 
i-o 

I'D 


range 

26-0-37-7 
31-0-39-6 
31-6-36-2 
31-4-37-7 
30-0-36-0 

26-0-39-6 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


n 

36 
10 

5 
7 
5 

63 


Head  length 
x  s.d. 


24-0 
24-7 

25*3 
24-9 

24-3 
24-4 


1-7 

i-o 
0-7 

i'3 

0-6 


s.e. 
0-3 
o-3 
o-3 
o-5 
o-3 
0-8 


range 

21-4-30-6 
23-6-26-9 
24-4-26-3 
23-1-26-2 
24-0-25-3 

21-4-30-6 


K.   E.   BANISTER 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


n 

36 
10 

5 

7 

4 

62 


Eye  diameter 
x 

7-0 
6-7 

7-2 

5-7 
7-0 

7-0 


s.d. 

i-i 

0-7 


s.e. 

0-2 
0-4 

O-2 
0-2 

0-2 


range 
4-3-10-2 

5-7-  8-3 
6-9-  8-4 
5-3-  6-3 
5-5-  7'3 
4-3-10-2 


Locality 

n 

X 

Nile 

36 

8-1 

Lake  Albert 

10 

8-6 

Blue  Nile 

5 

8-2 

Lake  Abaya 

7 

8-4 

Lake  Rudolf 

4 

8-0 

Total 

62 

8-2 

Interorbital  width 
s.d. 

0-8 
i-o 
0-8 
0-4 
o-3 
0-8 


s.e. 
o-i 

o-3 
0-4 
o-i 

O-I 

o-i 


range 

7-0-11-7 
7-0-10-5 

7-2-  8-7 
8-1-  8-8 
7'7-  8'3 
7-0-11-7 


Mouth  width 

Locality 

n 

X 

s.d. 

s.e. 

Nile 

36 

5'5 

0-7 

o-i 

Lake  Albert 

10 

6-0 

0-6 

0-2 

Blue  Nile 

5 

5'9 

0-4 

O-2 

Lake  Abaya 

7 

6-2 

0-7 

o-3 

Lake  Rudolf 

4 

5-o 

0-7 

o-3 

Total 

62 

5'7 

0-7 

O-I 

range 

4-2-7-3 
4-9-6-9 

5-4-6-5 
5-4-6-9 

4-3-5-8 
4-2-7*3 


Pectoral  fin  length 


Locality 

n 

X 

s.d. 

s.e. 

range 

Nile 

36 

21-3 

i-3 

0-2 

18-4-24-3 

Lake  Albert 

10 

21-5 

i-o 

o-3 

19-0-22-6 

Blue  Nile 

5 

24-5 

i-3 

0-6 

23-1-25-8 

Lake  Abaya 

7 

24-2 

1-9 

0-7 

21-6-26-4 

Lake  Rudolf 

5 

21-0 

1-2 

o-5 

19-6-22-5 

Total 

63 

21*9 

1-8 

O-2 

18-4-26-4 

Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


36 
10 

5 

7 

4 

62 


Caudal  peduncle  length 
x  s.d. 


17-0 
16-4 
16-2 
17-1 
17-1 

16-9 


0-7 
1-8 
0-6 


s.e. 
0-2 

o-3 
0-7 


range 

13-5-21-1 
14-7-19-7 
15-4-17-0 
15-5-20-6 
16-6-17-5 

13-5-21-1 


A   REVISION   OF  THE   LARGE   AFRICAN    BARBUS 


Caudal  peduncle  depth 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


n 

36 
10 

5 

7 
5 

63 


n 

36 
10 

5 

7 
5 

63 


34 
10 

5 

7 

5 

61 


34 
10 

5 

7 

5 

61 


M'3 
13-5 
13-6 

13-4 
13-8 


s.d. 

1-6 
0-9 

o-5 
0-7 
o-i 


Snout  length 

x  s.d. 


7'7 
7-8 
8-1 
8-2 
8-0 


0-7 
0-9 

o-3 
0-4 

o-3 
0-7 


Anterior  barbel 


4-6 

2'4 
6-3 
5'7 
2-5 

4'4 


s.d. 

1-4 
0-8 
i-i 


1-4 


Posterior  barbel 
x  s.d. 


6-5 
6-1 
4-0 

5'5 


i-o 
i-i 


O-2 
1-2 


s.e. 
0-3 
o-3 

0-2 

o-3 
o-i 

0-2 


s.e. 

o-i 
0-3 
o-i 
o-i 
o-i 

O-I 


s.e. 

0-2 
o-i 
0-4 

0-2 
0-2 

O-I 


s.e. 
o-i 
o-3 

0-2 

o-i 
o-i 

o-i 


range 
11-7-19-4 

13-1-15-4 
12-6-18-0 
12-6-14-6 
13-0-13-9 

11-7-19-4 


range 

6-7-10-5 
5-7-  9-2 
7-7-  8-4 
7-6-  8-8 
7'5-  8-3 
5-7-IQ-5 


range 

2-3-6-0 
1-2-4-6 
5-7-6-9 
4-6-6-1 
2-1-2-9 

1-2-6-9 


range 

4-2-8-4 
3-2-6-9 
6-1-7-4 
5-7-6-8 
3-9-4-3 
3-2-8-4 


Of  particular  note  is  the  ratio  of  the  caudal  peduncle  length  to  its  depth,  which 
has  a  mean  value  of  1-23.  Only  one  specimen  that  I  examined  had  a  ratio  greater 
than  1-40. 

The  figures  presented  here  show  that  the  barbels  are  shorter  in  fishes  from  Lakes 
Rudolf  and  Albert  than  they  are  in  the  other  localities  but  the  significance,  consid- 
ering the  inequality  of  the  samples,  is  low. 

On  the  basis  of  the  evidence  from  morphometrics  any  attempt  to  maintain  or 
establish  sub-specific  categories  is  rather  unsound.  Any  such  attempts  must  be 
supported  by  evidence  not  available  to  the  museum  worker. 

Squamation.  The  scales  of  Barbus  bynni  have  slightly  irregular  parallel  striae 
(fig.  93).  There  are  from  28  to  37  scales  in  the  lateral  line  series.  The  range  and 


K.    E.    BANISTER 


the  mean  (to  the  nearest  whole  number)  of  scales  on  the  lateral  lines  from  the 
different  localities  are  given  below. 


Locality 

Nile 

Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


33 

32 
3i 
32 
33 
33 


range 

31-37 
28-35 
28-33 
31-34 
28-35 

28-37 


Six  and  a  half  (less  frequently  5-5)  scales  are  present  between  the  dorsal  mid-line 
and  the  lateral  line.  Five  and  a  half  (rarely  6-5  or  4-5)  scales  are  present  between 
the  lateral  line  and  the  ventral  mid-line.  The  number  of  scales  between  the  lateral 
line  and  the  base  of  the  ventral  fin  varies  from  2  to  4-5 ;  then,  2  (f.i),  2-5  (f.2o),  3  (f.2o), 
3-5  (f.i4),  4  (f-7),  4-5  (f.i).  There  is  no  apparent  correlation  between  locality  and 
this  count.  There  are  from  12  to  16  scales  around  the  caudal  peduncle  ;  12  (f.8), 
13  (f.22),  14  (f.i7),  15  (1.7),  16  (f.7).  The  fish  from  the  Blue  Nile  had  12  scales 
around  the  caudal  peduncle  whilst  those  from  Lake  Rudolf  had  16.  Two  of  the 
Lake  Albert  fish  also  had  16.  The  samples  are  too  small  though  to  use  such  a 
variable  character  to  establish  a  population  taxonomically. 

Dorsal  fin.  The  dorsal  fin  has  8  (f.i),  9  (1.59),  10  (f.3)  branched  rays,  the  fourth 
unbranched  ray  is  ossified  into  a  straight,  massive  spine. 

Dorsal  spine. 


White  Nile 
Lake  Albert 
Blue  Nile 
Lake  Abaya 
Lake  Rudolf 

Total 


36 
10 

5 
7 
5 

63 


29-3 
30-9 
36-4 
35'3 


s.d. 

3'4 
2-5 
5-o 
3'9 
5'2 

4'3 


s.e. 

0-6 
0-8 

2-2 

i-5 
2-3 
0-6 


range 

23-7-38-4 
24-9-35-2 
29-5-40-0 

28-8-39-3 
30-0-43-0 

23-7-43-0 


The  anal  fin  has  three  unbranched  rays  and  five  branched  rays. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  teeth  (fig.  18)  number  five  in  the 
inner  row.  The  first  tooth  is  small  and  angled  towards  the  second  tooth.  This 
tooth  becomes  mammilliform  with  increasing  size  but  is  almost  always  higher  than 
it  is  long  (cf.  Barbus  gananensis).  Small  concavities  develop  on  the  posterior  face 
of  the  crown  on  teeth  3,  4  and  5.  These  three  teeth  become  successively  more 
slender  and  the  fifth  tooth  becomes  markedly  hooked  in  large  fishes.  The  three 
teeth  of  the  second  row  and  the  two  teeth  of  the  third  row  resemble  the  last  tooth 
in  the  inner  row  in  shape.  A  series  of  pharyngeal  bones  (fig.  19)  shows  the  variation 
in  shape  with  the  growth  of  the  fish. 

Gill  rakers.     There  are  12  to  1 6  gill  rakers  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  Described  by  Boulenger  (191  la)  as  yellowish  or  silvery,  grey  or  olive 
on  the  back,  fins  yellow  or  orange,  anal  and  caudal  pink  or  red,  ends  of  dorsal  and 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


33 


FIG.  1 8.     Dorsal  view  of  the  left  pharyngeal  of  -B.  bynni  of  280  mm  S.L. 

caudal  rays  sometimes  blackish.  Amirthalingam  (1965),  describing  Barbus  bynni 
from  the  Sudan,  states  that  the  colour  is  very  variable,  but  in  live  fish  of  about 
250  mm  is  frequently  as  follows  :  body  silvery,  dark  above  and  light  below  ;  all 
fins  with  deep-orange  streaks  ;  dorsal  and  caudal  fins  edged  with  black  ;  anal, 
pelvic  and  pectoral  fins  edged  with  white. 

DISTRIBUTION.  Specimens  from  Lake  Abaya,  Lake  Rudolf  and  Lake  Albert 
have  been  examined.  Sandon  (1950)  states  that  Barbus  bynni  is  common  in  the 
Blue  Nile  and  I  have  seen  specimens  from  the  Metti  and  Gudar  rivers  and  from 
Rosaires.  Sandon  (op.  cit.)  also  refers  to  the  widespread  distribution  of  Barbus  bynni 
throughout  the  White  Nile,  although  the  B.M.  (N.H.)  collections  have  no  specimens 
from  between  Khartoum  and  Lake  Albert.  This  species  is  common  throughout 
the  lower  Nile. 

Specimens  nos.  92-101  listed  by  Boulenger  (igua  :  28)  from  Lake  Baringo  are 
not  referable  to  this  species,  but  belong  to  Barbus  intermedius. 

Peel  (1900  :  336)  records  Barbus  byrni  (sic)  from  the  Webi  Shebeli  in  Somaliland. 
Donaldson-Smith  (1897)  collected  three  specimens  of  a  Barbus  species  identified  as 
Barbus  bynni  by  Giinther  (1896)  ;  the  two  small  specimens  from  Stony  Brook,  the 
Errer  river  (Webi  Shebeli  system)  (Academy  of  Natural  Sciences  Philadelphia,  Nos. 
14541,  14542),  are  not  Barbus  bynni  but  the  large  specimen  is.  The  two  small  fishes 
are  mentioned  with  details  of  the  locality  in  Donaldson-Smith's  account  of  his 
expedition  but  there  is  no  mention  of  the  large  fish.  Giinther  (1896)  gives  its 
locality  as  the  Shebeli  river.  This  specimen  is  now  in  the  collection  of  the  Philadel- 
phia Academy  of  Natural  Sciences  No.  16710,  where  the  locality  Thebeti,  East 
Africa,  has  been  given  to  it.  Thebeti  could  well  be  a  mis-reading  of  Shebeli,  for  the 


34 


K.   E.    BANISTER 


FIG.  19.  A  series  of  pharyngeal  bones  from  specimens  of  B.  bynni  to  show  the  variation 
with  the  size  of  the  fish.  Standard  lengths  (mm)  of  the  specimens  are  A  =  58  ;  B  =  98  ; 
C  =  140  ;  D  =  260  ;  E  =  235  ;  F  =  171  ;  G  =  280  ;  H  =  350. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


35 


^rr^A^Jo-/ 


FIG.  20.     Map  of  the  distribution  of  B.  bynni. 

writing  on  the  label  inside  the  gill  cover  is  very  faint.  This  fish  is  certainly  Barbus 
bynni  but  its  locality  is  in  some  doubt  and  it  is  not  included  with  the  data  presented 
above.  Its  measurements,  however,  will  appear  at  the  end  of  this  section. 

Vinciguerra  (1898)  gives  Bissan  Guarrica  as  a  locality  for  Barbus  ruspolii.  As 
far  as  I  can  find  out  this  is  a  tributary  of  the  Sagan  river  which  drains  into  Lake 
Stephanie. 

Lakes  Rudolf  and  Abaya  are  now  isolated  from  the  Nile  system.  Fryer  (1968) 
suggests  that  the  Southern  Ethiopian  rift  valley  lakes  drained  into  Lake  Rudolf  and 
Lake  Rudolf  was  connected  to  the  Nile  via  the  Sobat  river  (or  via  the  present 
Lotagipi  and  Kenamuki  swamps  as  suggested  by  Cooke  1958).  Therefore  Barbus 
bynni  could  have  been  isolated  in  the  distal  parts  of  what  was  once  a  more  extensive 
Nile  system.  Argulus  rhipidiophorus ,  a  parasitic  crustacean  discussed  by  Fryer 
(op.  cit.)  is  also  found  in  the  Nile,  Lake  Rudolf  and  the  Ethiopian  rift  valley  lakes. 
Greenwood  (1959)  has  recorded  remains  of  fishes  closely  resembling  Barbus  bynni 
from  an  epi-Pleistocene  site  at  Ishango  near  the  source  of  the  Semliki  river  on  Lake 
Edward.  Barbus  bynni  is  now  extinct  in  Lake  Edward  (see  p.  26). 


36  K.    E.    BANISTER 

Barbus  bynni  is  confined  to  the  Nile  system  and  is  also  found  in  lakes  that  were 
once  connected  to  the  Nile. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  bynni  has  some  gross  morphological  charac- 
ters in  common  with  Barbus  gananensis,  Barbus  oxyrhynchus  and  Barbus  longifilis. 
The  characters  in  common  include  a  rhomboidal  compressed  body,  a  strong,  straight, 
smooth  dorsal  spine  and  parallel,  sinuous  striae  on  the  scales.  The  species  listed 
above  are  considered  to  be  closely  related  and  comprise  the  Barbus  bynni  complex 
which  is  discussed  further  on  page  128. 

Barbus  bynni  can  be  distinguished  from  Barbus  gananensis  by  the  length  and 
strength  of  the  dorsal  spine  (longer  and  stronger  in  the  former  species)  and  by  the 
shape  of,  particularly,  the  second  tooth  in  the  inner  pharyngeal  row.  In  Barbus 
gananensis  this  tooth  is  higher  and  longer  than  in  Barbus  bynni  (see  fig.  28).  The 
anterior  edentulous  process  of  the  pharyngeal  bone  is  relatively  longer  in  Barbus 
bynni  than  in  Barbus  gananensis. 

The  other  two  species  are  well  separated  geographically  from  Barbus  bynni. 
Barbus  longifilis  can  easily  be  distinguished  by  its  barbels  (Ab,  x  =  8-9,  range 
8-3-9-6  ;  Pb,  x  =  ii'3,  range  9-5-13-4)  which  are  much  longer  than  those  of  Barbus 
bynni  (Ab,  x  =  4-4,  range  1-2-6-9  >  Pb,  x  =  5-5,  range  3-2-8-4). 

Barbus  oxyrhynchus  has  fewer  scales  in  the  lateral  line  series  than  has  Barbus  bynni 
(21-29,  most  frequently  22-25  against  28-37,  most  frequently  31-33).  The  dorsal 
spine  is  usually  thinner  and  shorter  in  Barbus  oxyrhynchus  than  in  Barbus  bynni 
(x  =  25-3  in  Barbus  oxyrhynchus,  x  ==  31-1  in  Barbus  bynni). 

Barbus  bynni  P.A.S.  No.  16710  (see  above) 

L  =  349  mm  ;  D  =  31-0  ;  H  =  21-8  ;  1  =  5-4;  IO  =  8-6  ;  MW  =  5-1  ;  Pet 
=  23-2  ;  CP1  =  14-6  ;  CPd  =  13-4  ;  Snt  =  6-9  ;  Ab  =  4-0  ;  Pb  =  4-9  ;  DSp 

=  29-3- 

Dorsal  fin  IV-g.  Twelve  scales  around  caudal  peduncle  ;  31  in  lateral  line, 
5-5  between  dorsal  mid-line  and  lateral  line,  5-5  from  lateral  line  to  ventral  mid- 
line  ;  2-5  between  lateral  line  and  base  of  pelvic  fin. 


Barbus  caudovittatus  Boulenger  1902 

Barbus  caudovittatus  Blgr.,  1902,  Annls.  Mus.  r.  Congo  Beige  Zool.  (i)  2  :  32  ;  Poll,  1953,  Result. 

scient.  Explor.  hydrobiol.  Lac  Tanganika  3  (5 A)  :  85. 
Varicorhinus  stappersii  Blgr.,  1917,  Ann.  Mag.  nat.  Hist.  (8)  20  :  364. 
Barbus  euchilus  Blgr.,  1919,  Proc.  zool.  Soc.  Lond.  :  400. 
Barbus  miochilus  Blgr.,  1919,  Proc.  zool.  Soc.  Lond.  :  401. 
Barbus  lestradei  David,  1936,  Revue  Zool.  Bot.  afr.  27  (2)  :  150,  fig.  i. 
Barbus  chilotes  sakaniae  Poll,  1938,  Revue  Zool.  Bot.  afr.  30  (3)  :  413,  fig.  13. 
Barbus  pojeri  Poll,  1944,  Bull.  Mus.  r.  Hist.  nat.  Belg.  20  (3)  :  2,  fig.  3. 

NOTES  ON  THE  SYNONYMY.  Poll  (1946)  synonymized  Barbus  euchilus  and  Barbus 
miochilus  and  then  in  1953  included  Barbus  pojeri  within  the  expanded  Barbus 
euchilus.  After  examination  of  the  type  specimens  I  agree  with  Poll,  although  it  is 
worth  noting  that  the  dorsal  fin  spines  of  Barbus  euchilus  and  Barbus  miochilus  are 
more  strongly  ossified  than  is  the  norm  in  Barbus  caudovittatus. 


A   REVISION   OF  THE  LARGE   AFRICAN   BARBUS  37 

Barbus  lestradei  had  been  recognized  as  a  separate  species  by  authors  since  its 
first  description  by  David  (1936) .  The  type  specimen  in  the  Central  African  Museum 
at  Tervuren  has  IV-g  rays  in  the  dorsal  fin  (not  II-Q)  and  27  scales  in  the  lateral 
line.  It  is  a  large  fish  (S.L.  373  mm)  as  are  almost  all  fish  referred  to  Barbus  lestradei. 
The  body  is  dark  coloured,  but  the  measurements,  the  weakly  ossified  dorsal  spine, 
overall  body  shape  and  the  indications  of  darkening  in  body  colour  in  some  Barbus 
caudovittatus  leave  little  room  for  doubt  that  Barbus  lestradei  is  a  synonym  of 
Barbus  caudovittatus.  Varicorhinus  stappersii  was  described  from  two  specimens  of 
255  mm  and  191  mm  S.L.  respectively  from  Nganza  on  the  Lufuko  river,  Lake 
Tanganyika  basin  B.M.  (N.H.)  Reg.  No.  1920.5.25  :  36-37,  and  two  specimens  from 
the  same  locality,  M.A.C.T.  No.  14222  of  S.L.  175  and  184  mm.  The  morphometric 
data  for  the  two  B.M.  (N.H.)  specimens  (not  included  below)  are  D  =  29-0,  29-3  ; 
H  =  20-8,  22-5  ;  I  =  47,  5-8  ;  IO  =  8-8,  8-6  ;  MW  =  9-4,  8-6  ;  Pet  =  21-2, 
20-9  ;  CP1  =  19-2,  19-4  ;  CPd  =  n-8,  11-5  ;  Snt  =  7-5,  8-1 ;  Ab  =  3-9,  4-7  ; 
Pb  =  5-5,  6-5  ;  DSp  =  9-4,  io-o  ;  D.  fin  IV-io,  IV-g.  The  morphometric  data 
of  these  two  fish  are  given  separately  to  facilitate  a  comparison  with  the  morpho- 
metric data  of  the  other  specimens  of  Barbus  caudovittatus.  There  are  12  scales 
around  the  caudal  peduncle,  28  in  the  lateral  line,  4-5,  5-5  in  the  transverse  series 
and  2-5  between  the  lateral  line  and  base  of  the  pelvic  fin.  The  M.A.C.T.  specimens 
agree  closely  with  these  measurements. 

The  mouth  is  broad  with  a  straight,  horny  edge  to  the  lower  jaw.  The  width  of 
the  mouth  is  a  natural  consequence  of  achieving  the  'Varicorhinus'  facies,  as  per- 
haps is  the  decrease  in  head  length.  The  barbels  are  conspicuous  and  the  pharyngeal 
bones  are  identical  with  those  of  Barbus  caudovittatus.  These  features  and  the 
similarity  between  the  morphometric  data  of  Varicorhinus  stappersii  and  Barbus 
caudovittatus  suggest  that  the  two  specimens  of  Varicorhinus  stappersii  are  highly 
modified  'sector'  forms  of  Barbus  caudovittatus  (Groenewald  1958).  The  situation 
here  parallels  the  case  of  Varicorhinus  brucii  and  Barbus  brucii,  two  species,  display- 
ing different  facies,  that  were  eventually  synonymized  by  Jubb  (1968).  It  is  not 
within  the  scope  of  this  paper  to  discuss  the  validity  of  the  genus  Varicorhinus,  but 
there  is  sufficient  awareness  of  the  variability  of  Barbus  species  mouth  parts  to 
suggest  that  it  is  not  unlikely  that  some  African  Varicorhinus  species  are  highly 
modified  individuals  of  various  polymorphic  Barbus  species.  The  specimens  from 
Koki,  recorded  by  Poll  (1953),  are  only  doubtfully  Barbus  caudovittatus.  The 
three  fishes  examined,  B.M.  (N.H.)  1955.12.20 : 870-872,  ex  Poll,  are  small, 
43-47  mm  S.L.  with  10  or  u  branched  rays  in  the  dorsal  fin,  a  well-ossified  dorsal 
spine,  24-25  scales  in  the  lateral  line  and  no  dark  marks  on  the  caudal  fin.  Although 
these  Koki  specimens  come  from  within  the  distributional  limits  of  Barbus  caudo- 
vittatus I  have  too  little  information  to  decide  whether  they  are  a  separate  species 
or  a  distinct  population  of  Barbus  caudovittatus.  They  have  not  been  considered  to 
be  Barbus  caudovittatus  in  this  account. 

HOLOTYPE.  A  fish  of  77  mm  S.L.,  M.A.C.T.  No.  1168,  from  Banzyville.  Another 
specimen  listed  as  'one  of  the  types'  is  in  the  collections  of  the  British  Museum 
(Natural  History)  No.  B.M.  (N.H.)  1901.12.26  :  26. 


K.   E.    BANISTER 


FIG.  21.     Barbus  caudovittatus  (Holotype,  from  Boulenger 


DESCRIPTION.     The  description  is  based  on  47  specimens, 
data  are  expressed  in  tabular  form  as  follows  : 


The  morphometric 


L 

D 

H 

I 

IO 

MW 

Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


28-8 

26-6 

7-5 

9-4 

7-3 

21-0 

16-8 

12-2 

8-9 
4'4 
5-8 


s.d. 
3'4 

2-0 

1-6 
1-6 

0-8 


1-4 
i-i 
i-i 
1-6 


s.e. 


O-2 
0-2 
0-1 
0-2 
O-2 
0-2 
0-2 
O-2 
O-2 


range 
35    -580  mm 

24-1-37-5 
21-4-31-4 

4-7-11-4 

6-5-12-4 

5-7-  8-9 
18-1-24-7 
12-8-19-7 

8-6-15-8 

6-5-11-4 

2'3-   7'5 

3-1-11-3 


Small  specimens  have  a  shallow  compressed  body,  but  this  becomes  deeper  and 
thicker  in  larger  fish.  The  mouth  is  small,  inferior  and  frequently  horse-shoe 
shaped.  Thicker  lips  have  been  observed  in  some  larger  specimens  from  Lake 
Tanganyika,  whilst  in  a  specimen  from  the  Luilu  river  (Sankuru  system  -  Kasai  ; 
M.A.C.T.  No.  78925),  the  'rubber-lipped'  condition  is  approached. 

The  ventral  profile  of  the  body,  from  the  jaw  articulation  to  the  anus,  is  straight 
or  slightly  convex.  The  illustrated  specimen  (the  holotype)  is  not  typical  in  this 
respect.  The  contrast  between  the  ventral  and  dorsal  profiles  is  usually  greater  in 
larger  fishes. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays.  In  one  fish  from  the 
Luberizi  river  (Ruzizi  area)  there  are  only  three  unbranched  rays.  David  (1936) 
recorded  only  two  unbranched  rays  in  the  type  specimen  of  Barbus  lestradei,  but  on 
examination  of  the  type  I  found  that  there  are  four.  The  last  unbranched  ray  is 
scarcely  ossified  and  in  the  majority  of  specimens  articulations  persist  almost  to  its 
base.  A  specimen  from  Nyundeulu  (an  affluent  of  the  Lualaba),  M.A.C.T.  No. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


39 


FIG.  22. 


1cm 


Dorsal  and  lateral  views  of  the  right  pharyngeal  bone  from  a  large 
specimen  of  B.  caudovittatus  of  420  mm  S.L. 


124937,  has  an  ossified  fourth  dorsal  ray  with  a  length  of  17  per  cent  of  the  standard 
length. 

The  branched  rays  range  from  8  to  u  in  number  ;  8  (f.3),  9  (f-40),  10  (f.3), 
ii  (f.i).  The  fishes  with  10  and  n  branched  rays  in  the  dorsal  fin  all  come  from 
the  river  at  Elizabeth ville  (=  Lubumbashi).  The  last  branched  ray  is  commonly 
longer  than  the  few  rays  immediately  preceding  it. 

The  anterior  edge  of  the  dorsal  fin  is  in  advance  of  the  insertion  of  the  pelvic  fin. 
Because  of  the  softness  of  the  last  unbranched  ray  it  was  impractical  to  measure  its 
length  using  the  same  standards  as  for  the  other  species. 

The  anal  fin  has  three  unbranched  and  five  branched  rays. 

Squamation.  The  lateral  line  has  24-30  scales,  24  (f.3),  25  (f.6),  26  (f.12),  27  (f.i3), 
28  (f.8),  29  (f.4),  30  (f.i).  There  is  no  evidence  for  any  geographical  variation  in 
these  figures.  There  are  12  scales  round  the  caudal  peduncle  and  4-5  (r.  5-5)  scale 
rows  between  the  dorsal  mid-line  and  the  lateral  line  and  4-5  (r.  5-5)  between  the 
lateral  line  and  the  ventral  mid-line.  Two  and  a  half  (r.  3)  scale  rows  are  present 
between  the  lateral  line  and  the  base  of  the  pelvic  fin. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  bones  of  a  large  specimen  are  shown 
in  fig.  22  and  of  a  small  specimen  in  fig.  23. 

The  crowns  of  the  teeth  are  hooked  in  small  fishes  but  become  molariform  in  large 
fish.  No  exception  to  the  pharyngeal  formula  2. 3. 5. -5. 3. 2  was  found. 

Gill  rakers.  The  number  of  gill  rakers  on  the  lower  limb  of  the  first  gill  arch  is 
between  11  and  15  in  the  specimens  examined. 


40  K.    E.    BANISTER 


FIG.  23.     Left  pharyngeal  bone  from  a  specimen  of  B.  caudovittatus  of  no  mm  S.L. 


Coloration.  In  small  specimens  (less  than  about  50  mm  S.L.)  a  small  black  spot 
is  present  on  the  caudal  peduncle  at  the  base  of  the  caudal  fin.  This  spot  is  not 
present  on  larger  specimens  but  instead  the  dorsal  and  ventral  edges  of  the  caudal 
fin  become  darker.  In  fishes  of  from  about  70  to  170  mm  S.L.  the  dark  bands  on 
the  caudal  fin  are  conspicuous.  Above  this  size  the  bands  are  less  obvious  as  a 
result  of  the  general  darkening  of  the  caudal  fin. 

In  most  specimens  dark  pigment  is  present  on  the  distal  parts  of  the  anal,  pelvic 
and  pectoral  fins.  This  is  most  noticeable  in  medium-sized  fishes. 

The  body  is  brown  in  alcohol-preserved  specimens.  Large  fishes  are  darker 
than  small  fishes. 

DISTRIBUTION.  Specimens  have  been  examined  fom  Elizabethville  (=  Lubum- 
bashi)  (Lualaba  Congo)  ;  Banzyville  ( =  Mobayi)  (Ubangi  Congo)  ;  Avakubi  (Ituri)  ; 
Tshikapa,  Luembe,  and  Tchitatu  rivers  (upper  Kasai)  ;  Bushiame  and  Luilu  rivers 
(Sankuru  system)  ;  Dundo,  Angola  ;  Stanleyville  ( =  Kisangani)  (Wamba  river)  ; 
Yangambe  (Isalowe  river  -  Uele  system)  ;  Luberizi  river  (Ruzizi  river)  and 
from  Lake  Tanganyika. 

Poll  (1953)  thinks  that  the  presence  of  Barbus  caudovitattus  (as  defined  here)  in 
Lake  Tanganyika  is  fortuitous  and  that  it  essentially  is  a  fluviatile  species. 

DIAGNOSIS.  Barbus  caudovittatus  is  a  slender-bodied  fish  characterized  by  a  very 
soft,  poorly  ossified  last  simple  ray  in  the  dorsal  fin  and  a  dark  band  on  each  lobe 
of  the  caudal  fin.  It  is  most  likely  to  be  confused  with  Barbus  trachypterus  and 
Barbus  paucisquamatus  which  it  superficially  resembles  in  body  shape,  scale  counts, 
and  mouth  position  and  mouth  shape.  It  is  not  yet  possible  to  determine  whether 
these  three  species  are  related  as  I  have  seen  too  few  specimens  of  the  latter  two 
species.  The  dorsal  spines  of  Barbus  trachypterus  (x  =  18-6,  range  16-1-21-3)  and 
Barbus  paucisquamatus  (x  =  19-7,  range  14-1-22-5)  are  much  longer  and  better 
ossified  than  is  normal  in  Barbus  caudovittatus,  but  a  few  specimens  have  developed 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  24.     Map  of  the  distribution  of  B.  caudovittatus. 


spines  of  up  to  17  per  cent  S.L.  Dark  pigment  is  present  on  the  distal  parts  of  the 
pectoral,  pelvic  and  anal  fins  in  Barbus  caudovittatus  and  Barbus  paucisquamatus 
but  on  the  proximal  parts  of  these  fins  in  Barbus  trachypterus.  The  barbels  are  much 
shorter  in  Barbus  caudovittatus  (Ab,  x  =  4-4,  range  2-3-7-5  ;  Pb,  x  =  5-8,  range 
3-1-11-3)  than  in  Barbus  paucisquamatus  (Ab,  x  =  7-5,  range  5-8-10-6  ;  Pb, 
x  =  8-6,  range  6-2-11-6). 


Barbus  ethiopicus  Zolezzi  1939 

Barbus  ethiopicus  Zolezzi,  1939,  Boll.  Pesca  Piscicolt.  Idrobiol.  Roma  15  :  369,  fig.  i. 

HOLOTYPE.  A  fish  of  circa  230  mm  S.L.  from  Lake  Zwai.  This  specimen  has 
not  been  seen  by  me  but  is  the  only  one  described  by  Zolezzi.  The  specimen  is 
housed  in  the  Laboratorio  centrale  di  Idrobiologia  in  Rome. 

DESCRIPTION.  The  description  is  based  on  three  specimens  of  240,  250  and 
258  mm  collected  in  Lake  Zwai  by  Mr  Eric  Hamblyn.  B.M.  (N.H.)  Nos.  1971.7.12  : 
1-3- 


K.   E.    BANISTER 


'I- 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  43 

The  range  of  ratios  (expressed  as  percentage  of  S.L.)  for  the  three  specimens  is 
as  follows. 

D  23-2-24-6  CP1  15-8-17-0 

H  25-8-27-1  CPd  8-4-10-0 

I  5-8-  6-2  Snt  8-8-  9-4 

IO  8-0-  8-8  Ab  3-6-  4-2 

MW  7-0-  7-9  Pb  3-6-  4-7 

Pet  16-3-18-7 

Calculation  of  the  standard  deviation  and  the  standard  error  with  only  three 
specimens  was  not  considered  a  profitable  action. 

The  elongate  body  is  round  in  cross-section  and  the  caudal  peduncle  is  long  in 
relation  to  its  depth.  The  mouth  is  wide  and  the  gape  is  terminal.  The  body  was 
covered  with  a  thick  layer  of  mucus  which  had  to  be  removed  before  scale  counts 
could  be  taken.  There  are  39  (=  18  +  21)  vertebrae. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays.  The  last  unbranched  ray 
is  weakly  ossified  as  a  smooth  spine  from  8-4  to  9-6  per  cent  of  the  standard  length. 
The  three  specimens  have  6  (f.i),  7  (f.i)  or  8  (f.i)  branched  dorsal  fin  rays.  The 
holotype  has  eight.  This  degree  of  variation  is  unusual  and  the  number  of  branched 
rays  is  low  for  the  large  Barbus  species  of  East  Africa  although  not  infrequent  in 
the  small  Barbus  species  with  radiately  striated  scales.  The  dorsal  fin  is  inserted 
slightly  in  front  of  the  pelvic  fins. 

The  anal  Jin  has  three  unbranched  and  five  branched  rays. 

Squamation.  The  lateral  line  has  46  (f.i),  48  (f.i)  or  52  scales  (48  and  50  on 
either  side  of  the  holotype),  a  number  much  higher  than  in  any  other  large  Barbus 
species  of  the  region.  There  are  18  scales  round  the  caudal  peduncle  (18-20  in  the 
holotype  according  to  Zolezzi).  Between  the  dorsal  mid-line  and  the  lateral  line 
there  are  9-5  (f.i)  or  10-5  (f.2)  scale  rows  and  10-5  (f.2)  or  11-5  (f.i)  between  the 
lateral  line  and  the  ventral  mid-line.  Five  and  a  half  scale  rows  are  present  between 
the  lateral  line  and  the  base  of  the  pelvic  fin. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  teeth  number  2. 3. 5. -5. 3. 2  (fig.  26), 
the  first  two  teeth  in  the  inner  row  are  conical  with  small  depressions  on  the  posterior 
face  of  the  crown.  In  succeeding  teeth  the  concavity  becomes  larger  and  as  the 
stem  of  the  tooth  becomes  thinner  the  whole  tooth  assumes  a  spatulate  appearance 
with  a  recurved  tip. 

There  are  10  (f.i)  or  n  (f.2)  gill  rakers  on  the  lower  limb  of  the  first  gill  arch  in  the 
three  specimens  examined. 

Coloration.  The  preserved  fishes  are  olive  grey-brown  on  the  back  and  flanks  to 
just  below  the  lateral  line.  The  ventral  part  of  the  flanks  and  the  belly  are  orangeish 
to  pinkish-brown.  The  dorsal  fin  and  the  dorsal  lobe  of  the  caudal  fin  are  olive- 
brown.  The  other  fins  are  pale  brown. 

DISTRIBUTION.     Known  only  from  Lake  Zwai,  Ethiopia. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  ethiopicus  is  easily  distinguished  from  the 
other  species  described  in  this  paper.  It  has  a  shallow,  almost  cylindrical  body 


44  K.   E.   BANISTER 


FIG.  26.     Left  pharyngeal  bone  of  Barbus  ethiopicus.     Dorsal  and  lateral  views. 

with  more  than  46  scales  in  the  lateral  line.  Barbus  microterolepis ,  known  by  one 
specimen  from  a  stream  flowing  into  Lake  Zwai,  has  only  40  scales  and  a  much  more 
compressed  body. 

Small-scaled  Barbus  species  from  the  region  under  discussion  in  this  paper  are 
confined  to  lake  basins,  e.g.  Barbus  tropidolepis  and  Barbus  platyrhinus  of  Lake 
Tanganyika.  However,  it  is  not  possible  to  decide,  in  these  cases,  whether  the 
small  scales  are  an  adaptation  to  a  primarily  lacustrine  environment  or  not.  The 
geological  history  of  the  Lake  Zwai  basin  is  not  known  in  enough  detail  to  determine 
how  long  the  lake  was  isolated  from  the  major  rivers  of  the  area  and  hence  whether 
Barbus  ethiopicus  represents  a  relict  population  from  a  once  widespread  small- 
scaled  species  (and  here  it  must  be  noted  that  there  are  fluviatile  small-scaled 
species  in  South  Africa)  or  whether  it  is  a  modification  of,  for  example,  the  Barbus 
intermedius  stock  with  which  it  is  now  sympatric.  Barbus  intermedius  as  well  as 
Barbus  microterolepis  is  found  in  Lake  Zwai.  The  possible  relationships  of  the  latter 
species  to  Barbus  ethiopicus  are  discussed  on  p.  91. 

The  evidence  for  the  affinities  of  Barbus  ethiopicus  is  inconclusive.  There  are  no 
characters  which  preclude  its  descent  from  a  Barbus  intermedius  stock  but  equally 
there  are  no  characters  which  positively  suggest  a  relationship  with  such  a  stock. 

In  the  absence  of  positive  information,  the  affinities  of  Barbus  ethiopicus  are  left  in 
abeyance  and  it  is  not,  in  this  paper,  considered  to  be  related  to  Barbus  intermedius. 

Barbus  gananensis  Vinciguerra  1895 

Barbus  gananensis  Vinciguerra,   1895,  Annali  Mus.  civ.  Stor.  nat.  Giacomo  Doria  (2)   15  :  53, 
pi.  5,  fig.  2  ;    Vinciguerra,  1897,  Annali  Mus.  civ.  Stor.  nat.  Giacomo  Doria  (2)  17  :  362. 

HOLOTYPE.  The  holotype  M.S.N.G.  (Museo  Storia  Naturale  Genova)  No.  17525 
from  the  River  Ganana  (=  Juba)  between  Lugh  and  Bardera  in  Somaliland  was 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS 


45 


^^TO^^^g^^. 


FIG.  27.     Barbus  gananensis  (the  holotype  after  Vinciguerra  -  in  Boulenger  191  la). 

figured  by  Vinciguerra.  It  is  now  in  extremely  poor  condition  and  I  have  not  been 
able  to  examine  it  although  it  was  examined  for  me  by  Dr  E.  Tortonese.  I  have 
though,  through  the  kindness  of  Dr  E.  Tortonese,  been  able  to  examine  two  paralecto- 
types.  These  are  small  fish  of  50  and  39  mm  S.L.  and  they  came  from  the  Auata 
river  (=  Awata).  A  further  specimen  from  the  type  locality,  a  fish  of  176  mm  S.L., 
M.S.N.G.  No.  17339,  was  studied  and  this  specimen  is  illustrated  (fig.  28)  to  compare 
with  the  original  rather  poor  figure.  A  fourth  fish,  identified  as  Barbus  gananensis 
from  the  Citerni  collection  from  the  Upper  Ganana,  M.S.N.G.  No.  17343,  is  not 
referable  to  this  species. 

The  description  is  therefore  based  upon  three  specimens. 

DESCRIPTION.  As  a  result  of  the  description  being  based  on  two  small  fish  and 
one  larger  specimen,  certain  morphometric  data  show  a  large  range.  Only  the 
range  is  shown  as  other  calculations  could  be  misleading. 


L 

D 

H 

I 

10 

MW 


39    -176  mm 

28-2-32-0 

25-0-28-2 

7-4-10-2 

6-0-  8-5 

4'5-  7'7 


Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


23-1-24-0 

16-5-18-0 

13-1-15-0 

7-7-  8-0 

2-6-  6-3 

6-0-  7-9 


The  calculations  are  expressed  as  percentages  of  the  standard  length. 

The  body  is  compressed  and  similar  in  outline  to  the  body  of  Barbus  bynni, 
although  rather  less  deep.  The  caudal  peduncle  is  not  so  square  as  in  Barbus  bynni, 
the  ratio  of  the  CPl/CPd  is  from  1-2  to  1-3  in  Barbus  gananensis  against  i-o  to  1-2 
in  Barbus  bynni  of  the  same  size. 

Squamation.  The  scales  bear  parallel  striae,  slightly  less  sinuous  than  is  typical 
for  Barbus  bynni.  There  are  29  (f.2)  or  31  (f.i)  scales  in  the  lateral  line  and  12  scales 


K.    E.    BANISTER 


FIG.  28.     Barbus  gananensis,  a  specimen  of  176  mm  S.L. 


around  the  caudal  peduncle.  The  other  scale  counts  were  only  possible  for  the 
largest  specimen  and  here  there  are  5-5  scales  between  the  dorsal  mid-line  and  the 
lateral  line,  5-5  scales  between  the  lateral  line  and  the  ventral  mid-line  and  2-5 
scales  between  the  lateral  line  and  the  insertion  of  the  pelvic  fin. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays  and  nine  branched  rays. 
The  base  of  the  fin  is  enveloped  by  a  sheath  of  scales.  The  last  unbranched  ray 
forms  a  smooth  ossified  spine  from  18-0  to  22-7  per  cent  of  the  standard  length. 
This  is  rather  less  than  in  similar-sized  specimens  of  Barbus  bynni.  The  spine  in 
the  small  specimens  is  conspicuously  less  well  ossified  than  equal-sized  Barbus  bynni 
specimens.  The  origin  of  the  dorsal  fin  is  slightly  in  advance  of  the  origin  of  the 
pelvic  fin. 


FIG.  29.  A  comparison  of  the  left  pharyngeal  bones  from  a  specimen  of  B.  gananensis 
of  176  mm  S.L.  (left  bone)  with  the  left  pharyngeal  bone  from  a  Barbus  bynni  of 
171  mm  S.L. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  47 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays.  The  last 
branched  ray  is  markedly  bifurcated. 

Pharyngeal  bones  and  teeth.  The  contrast  between  the  pharyngeal  bones  of  the 
176  mm  S.L.  Barbus  gananensis  and  a  171  mm  S.L.  Barbus  bynni  can  be  seen  in 
fig.  29.  The  bone  is  thicker  and  deeper  in  the  former  and  the  teeth  of  the  inner  row 
are  higher.  The  second  tooth  of  the  inner  row  in  particular  is  much  longer  in  Barbus 
gananensis  than  in  Barbus  bynni.  These  differences  are  equally  clear  in  the  pharyn- 
geal bones  of  the  smaller  Barbus  gananensis  when  compared  with  equal-sized 
Barbus  bynni.  The  teeth  of  the  second  and  third  rows  are  only  marginally  more 
robust  than  in  comparable  Barbus  bynni  material. 

Gill  rakers.  There  are  12  gill  rakers  on  the  lower  limb  of  the  first  gill  arch  in  the 
largest  specimen  ;  the  gill  rakers  could  not  be  counted  in  the  smaller  specimens. 

Coloration.  In  alcohol,  the  young  fish  are  silvery  flanked,  with  a  brown  back. 
The  largest  fish  is  sandy  brown  on  the  flanks  and  belly,  darker  on  the  back. 

DISTRIBUTION.  Collected  from  the  Juba  river  (between  Lugh  and  Bardera)  and 
from  the  Awata  river  (upper  Juba).  Also  it  is  reported  from  the  confluence  of  the 
Ganana  Dulei  and  Bas  Nabor.  It  has  not  been  recorded  from  the  Webi  Shebeli 
where  Barbus  bynni  is  found. 

AFFINITIES.  A  species  that  is  very  close  to  Barbus  bynni,  differing  in  certain 
features  mentioned  above.  A  larger  collection  of  Barbus  gananensis  may  show 
modal  differences  further  distinguishing  it  from  Barbus  bynni.  The  general  im- 
pression culled  from  these  three  fish  is  that  they  resemble  a  Barbus  bynni  stock 
which  has  begun  to  approach  the  facies  of  the  Barbus  oxyrhynchus  from  the  Athi  and 
Tana  systems  immediately  to  the  south,  particularly  in  possessing  slightly  longer 
pectoral  fins  than  is  typical  for  Barbus  bynni  at  these  sizes,  a  reduced  dorsal  fin  spine 
and  no  less  than  nine  branched  rays  in  the  dorsal  fin.  However,  this  is  based  solely 
on  an  extremely  small  sample  of  fish  and  this  trend  (if  it  exists)  must  remain  hypo- 
thetical until  more  information  is  available  about  Barbus  gananensis. 


Barbus  intermedius  Riippell  1837 

Barbus  surkis  Riippell,  1837,  Mus.  senckenb.  2  :  5,  pi.  i,  fig.  if  ;    Blgr.,  1902,  Ann,  Mag.  nat. 

Hist.  (7)  10  :  424  ;    Blgr.,  1907,  Fish  Nile  :  226,  pi.  42  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  65. 
Barbus  intermedius  Rupp.,  1837,  Mus.  senckenb.  2  :  7,  pi.  i,  fig.  2  ;    Giinther,  1864,  Proc.  zool. 

Soc.  Lond.  :  91   (non  B.  intermedius  Riipp.  -fide  Blgr.,  1911  :  45)  ;    Blgr.,  1902,  Ann.  Mag. 

nat.  Hist.  (7)   10:424;    Blgr.,   1907,  Fish  Nile  :  228,  pi.  29,  fig.  2;    Blgr.,   1911,  Cat.  Afr. 

Fish  2  :  59,  fig.  39. 

f  In  Tracts  on  Natural  History,  vol.  4,  in  the  Zoological  Department  library  of  the  British  Museum 
(Natural  History)  (Cat.  No.  5  q  T  r)  there  is  a  separate  of  this  Riippell  paper  bearing  the  date  1835. 
The  title  page,  pagination  and  figures  are  identical  with  those  in  the  1837  Mus.  senckenb.  except  that  the 
title  page  bears  the  inscription  'Ausgezogen  aus  dem  II  Band  des  Museum  Senckenbergianum.  Frankfurt 
am  Main,  1835'. 

As  the  cover  of  the  second  volume  of  the  Mus.  senckenb.  bears  the  date  1836  and  the  title  page  the 
date  1837,  I  am  assuming  that  the  1835  date  is  either  a  typographical  error  or  an  over-optimistic  estimate 
of  the  date  of  publication.  Until  firm  evidence  to  the  contrary  is  available  I  shall  continue  to  refer  to 
Riippell  1837. 


48  K.   E.    BANISTER 

Barbus  intermedius  leptosoma  :    Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  167,  pi.  6,  fig.  i 

(vide  B.  leptosoma  Blgr.,  1902). 

Barbus  intermedius  microstoma  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  168,  pi.  6,  fig.  2. 
Barbus   intermedius   gorgorensis   Bini,    1940,    Missione   Stud.    Lago   Tana,   3    (2)  :  168,   pi.    6, 

fig-  3- 
Barbus  affinis  Riipp.,  1837,  Mus.  senckenb.  2  :  8,  pi.  i,  fig.  3  ;    Blgr.,  1902,  Ann  Mag.  nat.  Hist. 

(7)  10  :  425  ;    Blgr.,  1907,  Fish  Nile  :  216,  pi.  29,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  47, 

fig.  28. 
Barbus  affinis  brevibarbis  :    Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  162,  pi.  i,  fig.  2  (vide 

B.  brevibarbis  Blgr.,    1902). 
Barbus  affinis  nedgia  :    Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  163,  pi.  2,  figs,  i  and  2 

(vide  Labeobarbus  nedgia  Riipp.,  1837). 
Barbus gorguari i  Riipp.,  1837,  Mus.  senckenb.  2  :  9,  pi.  i,  fig.  4  ;  Blgr.,  1902,  Ann.  Mag.  nat.  Hist. 

(7)  10  :  424  ;    Blgr.,  1907,  Fish  Nile  :  237,  pi.  44  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  75,  fig.  53. 
Barbus  gorguarii  macrophtalmus  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  177,  pi.  12,  fig.  2. 
Barbus  elongatus  Riipp.,  1837,  Mus.  senckenb.  2  :  n,  pi.  2,  fig.  i. 
Labeobarbus  nedgia  Riipp.,  1837,  Mus.  senckenb.  2  :  14,  pi.  2,  fig.  3. 
Barbus  nedgia  (Riippell)  :    Gunther,  1868,  Cat.  Fish  7  :  104  ;    Blgr.,  1902,  Ann.  Mag.  nat.  Hist. 

(7)  10  :  426  ;    Blgr.,  1907,  Fish  Nile  :  223,  pi.  41,  fig.  2  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  51, 

fig-  32- 

Barbus  gregorii  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  422  ;   Blgr.,  1911,  Cat.  Afr.  Fish  2  :  45. 
Barbus  rueppelli  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  427  ;   Blgr.,  1911,  Cat.  Afr.  Fish  2  :  26, 

fig-  7- 
Barbus  macronema  Blgr.,    1902,   Ann.  Mag.  Nat.  Hist.   (7)   10:427;    Blgr.,   1911,   Cat.  Afr. 

Fish  2  :  25. 

Barbus  macronema  var.  parenzani  Zolezzi,  1940,  Boll.  Pesca.  Piscic.  Idrobiol.  15  :  372. 
Barbus  harringtoni  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  429  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2  :  61,  fig.  40. 
Barbus  plagiostomus  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  429  ;   Blgr.,  1907,  Fish  Nile  :  232, 

pi.  38,  fig.  2  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  68,  fig.  47. 
Barbus  jar  sinus  Blgr.,   1902,  Ann.  Mag.  nat.  Hist.  (7)   10:429;    Blgr.,   1911,  Cat.  Afr.  Fish 

2  :  62,  fig.  41. 
Barbus  platystomus  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  430  ;   Blgr.,  1907,  Fish  Nile  :  234, 

pi.  40,  fig.  2  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  73,  fig.  51. 

Barbus  platystomus  var.  vatovae  Zolezzi,  1939,  Boll.  Pesca.  Piscic.  Idrobiol.  15  :  370. 
Barbus  platystomus  platystomus  :   Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  170,  pi.  8,  fig.  i. 
Barbus  platystomus  daga  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  171,  pi.  8,  fig.  2. 
Barbus  platystomus  dekkensis  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  172,  pi.  9,  fig.  i. 
Barbus  platystomus  prognathus  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  173,  pi.  9,  fig.  2. 
Barbus  brevibarbis  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  431  ;    Blgr.,  1907,  Fish  Nile  :  219, 

pi.  35,  fig.  2  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  49,  fig.  29. 
Barbus  kassamensis  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  431  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2:  30,  fig.  ii. 
Barbus  oreas  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  432  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  38, 

fig.  19  ;   Pietschmann,  1913,  Jb.  Ver.  Naturk.  Wiesbaden  66  :  189. 
Barbus  leptosoma  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  432  ;    Blgr.,  1907,  Fish  Nile  :  220, 

pi.  40,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  50,  fig.  30. 
Barbus  duchesnii  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  433  ;    Pellegrin,  1905,  Bull.  Mus. 

Hist.  nat.  Paris  11  :  291  ;    Blgr.,    1907,  Fish  Nile  :  208,  pi.  35,   fig.   i  ;    Blgr.,    1911,   Cat. 

Afr.  Fish  2  :  31,  fig.  12. 

Barbus  duchesnii  maximus  Bini,   1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  165,  pi.  4,  fig.  2. 
Barbus  duchesnii  ibridus  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  166,  pi.  5. 
Barbus  mento  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  434  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  41, 

fig.  22. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  49 

Barbus  hursensis  Blgr.,  1902,  Ann.  Mag.  nat.  Hist.  (7)  10  :  434  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2  :  46,  fig.  27. 
Barbus  degeni  Blgr.,   1902,  Ann.  Mag.  nat.  Hist.   (7)   10  :  435  ;    Blgr.,   1907,  Fish  Nile  :  222, 

pi.  41,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  50,  fig.  31. 

Barbus  degeni  leptorhinus  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  164,  pi.  3,  fig.  2. 
Barbus  erlangeri  Blgr.,   1903,  Proc.  zool.  Soc.  Lond.   (2)  :  329,  pi.  29  ;    Blgr.,   1911,  Cat.  Afr. 

Fish  2  :  29,  fig.  10. 

Capoeta  bingeri  Pellegrin,  1905,  Bull.  Mus.  Hist.  nat.  Paris  11  :  292. 
Barbus  ilgi  Pellegrin,  1905,  Bull.  Mus.  Hist.  nat.  Paris  11  :  293. 
Barbus  neuvillii  Pellegrin,  1905,  Bull.  Mus.  Hist.  nat.  Paris  11  :  294. 
Barbus  zuaicus  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  562  ;   Blgr.,  1911,  Cat.  Afr.  Fish  2  :  64, 

fig.  43- 
Barbus  macmillani  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  562  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2  :  67,  fig.  46. 
Barbus  zaphiri  Blgr.,   1906,  Ann.  Mag.  nat.  Hist.  (7)   17  :  563  ;    Blgr.,   1907,  Fish  Nile  :  235, 

pi.  45,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  74,  fig.  52. 
Barbus  bottegi  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  563  ;   Blgr.,  1911,  Cat.  Afr.  Fish  2  :  70, 

fig.  48. 
Barbus  gudaricus  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  564  ;    Blgr.,  1907,  Fish  Nile  :  241, 

pi.  34,  fig.  2  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  40,  fig.  21. 
Barbus  eumystus  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  565  ;    Blgr.,  1907,  Fish  Nile  :  215, 

pi.  38,  fig.  i  ;    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  44,  fig.  25. 
Barbus  margaritae  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  565  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2  :  39,  fig.  20. 
Barbus  alticola  Blgr.,  1906,  Ann.  Mag.  nat.  Hist.  (7)  17  :  566  ;   Blgr.,  1911,  Cat.  Afr.  Fish  2  :  55, 

fig-  35- 

Barbus  bingeri  :    Blgr.,  1911,  Cat.  Afr.  Fish  2  :  69. 
Barbus  procatopus  Blgr.,  1916,  Ann.  Mag.  nat.  Hist.  (8)  17  :  244. 

Barbus  volpinii  Parenzan,  1940,  Boll.  Idrobiol.  Cacc.  Pesca.  Afr.  orient,  ital.  1  :  9,  fig.  i. 
Barbus  brunellii  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  173 
Barbus  brunellii  acutirostris  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  174. 
Barbus  dainellii  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  175. 
Barbus  dainellii  macrocephalus  Bini,  1940,  Missione  Stud.  Lago  Tana,  3  (2)  :  176. 
Varicorhinus  bingeri  :    Berlin  and  Esteve,  1947,  Catalogue  des  Types  des  Poissons,  Paris  4  :  44. 

NOTES  ON  THE  SYNONYMY.  Although  the  specific  name  surkis  appears  before 
intermedius  in  Riippell  (1837),  I  have  selected  the  latter  name  as  the  senior  synonym 
according  to  article  24  of  the  International  Code.  The  reasons  for  this  are  : 
a)  Ruppell's  text  states  that  Barbus  surkis  has  36  scales  in  the  lateral  line  but  his 
illustration  (pi.  i,  fig.  i)  shows  only  32  scales  ;  the  text  and  figure  of  Barbus  inter- 
medius agree  with  the  holotype  ;  b)  the  holotype  of  Barbus  surkis  is  untraceable 
but  the  holotype  of  Barbus  intermedius  exists  as  specimen  6778  in  the  Senckenberg 
Museum,  Frankfurt-am-Main  ;  c)  the  specific  name  intermedius  is  a  great  deal  more 
apposite  for  this  variable  species  than  is  surkis. 

A  synonymy  as  extensive  as  this,  involving  50  nominal  species  and  subspecies 
and  resulting  in  one  species  with  two  subspecies  requires  a  great  deal  of  justification. 
The  evidence  will  be  presented  below. 

Just  over  a  hundred  years  after  Riippell  described  the  deep-bodied  Barbus  surkis 
Bini  (1940)  described  the  elongated,  predatory-looking  Barbus  brunellii.  These 
two  species,  ironically  the  first  and  (to  date)  the  last  described  from  Lake  Tsana,  are 
very  different  in  shape  (see  figs.  31,  32  and  46)  and  a  simple  comparison  of  an 


5o  K.    E.    BANISTER 

example  of  each  form  would  undoubtedly  result  in  their  being  placed  in  separate 
species.  It  is  only  when  the  extremes  are  compared  with  other  Ethiopian  specimens 
that  it  becomes  evident  that  one  variable  species  is  present  instead  of  many,  scarcely 
distinct  species.  The  degree  of  variability  is  great  and  is  reflected  throughout  many 
parts  of  the  bodies  of  species  ;  head  length  may  be  singled  out  as  one  of  the  most 
variable  factors.  Because  of  this  plasticity  of  form  it  has  proved  impossible  to 
detect  more  than  two  statistically  recognizable  groups  within  the  whole  sample. 
It  had  been  suspected  by  Worthington  (ig32a)  that  this  would  be  the  case  from  the 
practical  difficulties  he  encountered  in  trying  to  distinguish  one  nominal  species 
from  another  using  the  published  descriptions. 

The  majority  of  species  were  described  between  1902  and  1906,  usually  from  a 
very  few,  often  poorly  preserved  specimens.  The  species  were  denned  on  the  basis 
of  small  differences  in  barbel  lengths,  lip  formation,  scale  counts  and  body  propor- 
tions ;  differences  that  may  be  tenable  with  few  specimens  but  lose  their  significance 
with  a  large  series.  Subsequent  workers  have  tried  with  increasing  difficulty  to  fit 
newly  acquired  specimens  into  the  framework  of  species  established  by  Boulenger 
(igna,  1916).  Bini  (1940)  reduced  several  of  Boulenger's  species  to  subspecific 
rank  but  at  the  same  time  described  a  further  two  species.  Worthington  (i932a  : 
127)  from  his  work  on  the  Kenyan  and  Ugandan  lakes  prophetically  commented, 
'  thus,  when  a  large  series  of  specimens  is  collected  from  Lake  Tsana  and  the  upper 
tributaries  of  the  Blue  Nile,  it  is  probable  that  the  series  of  species  endemic  to  the 
region  -  Barbus  leptosoma,  Barbus  degeni,  Barbus  nedgia  etc.,  will  prove  to  be  one 
and  the  same  species  since  they  differ  from  each  other  almost  solely  in  the  degree  of 
lip  development  and  in  the  size  of  the  eye  and  dorsal  spine'. 

The  graphs  and  calculations  presented  below  will,  I  hope,  show  the  essentially 
normal  distributions  of  the  characters  heretofore  considered  trenchant,  and  that  it  is 
biologically  more  reasonable  to  suggest  the  presence  of  one  variable  species  rather 
than  a  multitude  of  almost  indistinguishable,  ill-defined  or  indefinable  species. 

The  illustrations  reproduced  from  Boulenger  (igna)  and  Bini  (1940)  will  help  to 
supplement  the  calculations  and  show  the  various  intergrades  between  the  most 
extreme  forms. 

Worthington  (i932a  :  124)  noted  that  the  fishes  listed  by  Boulenger  (1911)  as 
Barbus  bynni  from  Lake  Baringo  should  be  placed  with  Barbus  gregorii  ( =  Barbus 
intermedius). 

Mann  (1971)  suggests  that  Barbus  gregorii  from  Lake  Baringo  is  rightfully  placed 
with  Barbus  tanensis  (=  Barbus  oxyrhynchus) .  His  comparative  material  was 
limited  to  small  fishes  (less  than  60  mm  S.L.)  which  can  give  rise  to  misleading 
conclusions  if  compared  with  larger  fish.  In  the  scale  counts,  nature  of  the  scale 
striae  and  in  most  morphometric  details  the  Lake  Baringo  fishes  are  identical  with 
Barbus  intermedius  from  Ethiopia.  However,  the  dorsal  spine  is  modally  longer 
than  in  the  other  Barbus  intermedius  populations  (although  the  spine  is  not  as  thick 
as  in  Barbus  tanensis}  and  the  pectoral  fin  is  longer.  Because  of  these  characters 
and  because  Lake  Baringo  is  isolated  and  the  most  southern  locality  for  this  species, 
I  think  it  would  be  valid  to  accord  sub-specific  status  to  the  Barbus  intermedius 
population  in  Lake  Baringo  and  I  propose  the  name  Barbus  intermedius  australis. 


A    REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  30.     Barbus  intermedius  (Holotype  from  Riippell  1837). 


FIG.  31.     Barbus  intermedius  (Holotype  of  B.  surkis  from  Riippell  1837). 

It  is  perhaps  worth  noting  that  the  longer  dorsal  spine  and  longer  pectoral  fin  which 
distinguish  the  Lake  Baringo  population  are  two  characters  possessed  in  greater 
degree  by  Barbus  oxyrhynchus. 

HOLOTYPE.  A  dried  specimen  of  296  mm  S.L.  No.  6778,  Senckenberg  Museum, 
Frankf  urt-am-Main . 

DESCRIPTION.  The  description  is  based  on  a  total  of  454  specimens  ;  179  from 
Lake  Tsana,  58  from  Lake  Baringo,  44  from  Lake  Abaya,  32  from  Lake  Zwai,  32 
from  the  Hawash  river,  30  from  Lake  Stephanie,  29  from  the  Blue  Nile,  24  from 


K.   E.    BANISTER 


FIG.  32.     Barbus  intermedius  (figure  of  B.  surkis  from  Boulenger  191  la). 


FIG.  33.     Barbus  intermedius  (Holotype  of  B.  dainellii  from  Bini  1940). 

Lake  Rudolf  and  the  Omo  river,  21  from  the  Webi  Shebeli  river,  four  from  Lake 
Langano  and  one  from  Lake  Orsodi. 

All  traceable  holotypes  have  been  examined  ;  those  not  seen  were  Barbus  surkis 
(Frankfurt,  missing  since  the  war),  Barbus  volpinii  (not  traceable),  Barbus  macronema 
var.  parenzani  (not  traceable)  and  Barbus  dainellii  (lost  through  poor  preservation) . 


n 

X 

s.d. 

s.e. 

range 

L 

454 

66    -489  mm 

D 

454 

28-4 

2-3 

o-i 

20-7-37-7 

H 

454 

26-4 

2-2 

O-I 

19-0-31-7 

I 

454 

5'9 

1-2 

0-05 

3-5-10-2 

10 

454 

8-0 

0-8 

0-03 

4-7-12-3 

MW 

450 

5'9 

I-O 

0-04 

3-6-  9-1 

Pet 

452 

21-8 

1-8 

o-i 

15-6-28-0 

CP1 

454 

16-9 

1-6 

o-i 

11-7-22-9 

CPd 

453 

n-6 

1-2 

o-i 

8-0-15-0 

Snt 

454 

8-5 

I-I 

0-05 

5-5-12-7 

Ab 

449 

5-8 

1-9 

o-i 

i-8-ii-i 

Pb 

45i 

6-9 

2-1 

o-i 

1-9-12-1 

A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  53 

B 


n  .  i  mi-*!:'!  ;  I 


5- 


imMmwn 


BN 


WS 


20      21       22      23      24      25      26      27      28      29      30      31      32      33      34      35 

FIG.  34.  Histograms  of  the  distribution  of  the  body  depth  (as  a  percentage  of  the  standard 
length)  in  B.  intermedius  populations  from  various  localities.  B  =  Lake  Baringo  ; 
O  =  Omo  river  ;  H  =  Hawash  river  ;  A  =  Lake  Abaya  ;  BN  =  Blue  Nile  ; 
T  =  Lake  Tsana  ;  WS  =  Webi  Shebeli  River  ;  Z  =  Lake  Zwai ;  S  =  Lake  Stephanie. 


54 


K.    E.    BANISTER 


70- 

60- 
50- 
40- 
30- 
20- 


10- 


TrmTmTTTnTm 


20   21   22   23   24   25   26   27   28   29   30   31   32   33   34   35 

FIG.  35.  Barbus  intermedius  :  a  composite  histogram  of  the  body  depth  for  specimens 
from  all  the  localities  treated  separately  in  Fig.  34.  The  distribution  of  body  depth 
throughout  the  whole  B.  intermedius  sample  is  normal. 


FIG.  36.     Barbus  intermedius  (Holotype  of  B.  degeni  from  Boulenger  191  la). 


FIG.  37.     Barbus  intermedius  (figured  specimen  of  B.  nedgia  in  Boulenger  1911  a). 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


55 


wwmmmtim 


FIG.  38.     Barbus  intermedius  (Holotype  of  B.  rueppelli  in  Boulenger  191  la). 


FIG.  39.     Barbus  intermedius  (figured  specimen  of  B.  gorguarii  from  Boulenger  191  la). 


The  morphometric  data  do  not  adequately  reflect  the  variation  in  body  form 
shown  by  this  species.  A  series  of  examples  of  different  facies  is  shown  in  figs.  36 
to  44.  Names  synonymized  above  are  used  below  for  convenience. 

The  body  varies  in  shape  from  the  slender  Barbus  dainellii  form  (fig.  33)  to  the 
deep  Barbus  surkis  form  (fig.  32).  It  can  be  seen,  though,  from  the  histograms  that 
in  the  whole  sample  the  body  depth  has  a  normal  distribution  (figs.  34,  35).  There 
is  a  little  difference  in  body  depth  distribution  in  different  localities,  e.g.  the  Lake 
Tsana  population  is  modally  less  than  the  Lake  Baringo  and  Omo  river  populations. 

The  mouth  and  lips  exhibit  a  considerable  degree  of  variation.  'Rubber-lipped' 
forms  are  common,  e.g.  the  forms  described  as  Barbus  degeni  (fig.  36)  and  Barbus 
nedgia  (fig.  37) .  Barbus  rueppelli  (fig.  38)  and  Barbus  gorguarii  (fig.  39)  have  large 
mouths  with  the  gape  at  45  degrees  to  the  horizontal ;  however,  Barbus  kassamensis 
(fig.  40)  and  Barbus  hursensis  (fig.  41)  link  this  mouth  type  with  the  ventral  or  sub- 
terminal  mouths  of  the  forms  referred  to  Barbus  oreas  (fig.  42)  and  Barbus  erlangeri 
(fig.  43).  The  ventral  mouth  with  a  horny  lower  lip  is  found  in  Barbus  macmillani 


56  K.   E.    BANISTER 

(fig.  44)  and  Barbus  bingeri.  Although  the  differences  seem  quite  distinct  when 
isolated  examples  are  considered,  examination  of  a  large  number  of  specimens  shows 
that  the  mouth  types  grade  into  each  other  without  any  sharp  divisions.  The  form 
of  the  mouth  in  this  species  is  valueless  as  a  taxonomic  character. 

Four  barbels  are  invariably  present  ;  the  Lake  Stephanie  population  has  modally 
slightly  longer  barbels  than  any  other  population,  but  this  is  very  possibly  a  reflec- 
tion of  the  size  range  of  the  sample.  The  barbels  are  relatively  smaller  in  larger 
fishes,  partly  because  of  negative  allometry  and  partly  because  of  the  increased 
chance  of  physical  damage.  Histograms  of  the  barbel  length  in  different  popu- 
lations are  shown  in  fig.  45.  The  head  length  varies  considerably.  The  forms  with 


FIG.  40.     Barbus  intermedius  (Holotype  of  B.  kassamensis  Boulenger  191  la). 


FIG.  41.     Barbus  intermedius  (Holotype  of  B.  hursensis  from  Boulenger  191  za). 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


57 


FIG.  42.     Barbus  intermedius  (Holotype  of  .B.  oreas  from  Boulenger  191  la). 


FIG.  43.     Barbus  intermedius  (Holotype  of  B.  erlangeri  from  Boulenger  191  la). 


FIG.  44.     Barbus  intermedius  (Holotype  of  B.  macmillani  from  Boulenger  191  la). 


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FIG.  45.  Histograms  of  the  distribution  of  the  anterior  and  posterior  barbel  lengths  in 
Barbus  intermedius  populations  from  various  localities.  The  lengths  are  expressed  as 
percentages  of  the  standard  length.  The  shaded  column  represents  the  anterior  barbel. 
The  locality  codes  are  as  in  Fig.  34. 


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FIG.  46.     Barbus  intermedius  (figured  examples  of  B.  brunelli  (above)  and 
B.  brunelli  acutirostris  from  Bini  1940). 

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FIG.  47.     Histograms  of  the  distribution  of  the  head  length  expressed  as  a  percentage 
of  the  standard  length  for  the  entire  Barbus  intermedius  sample. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


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FIG.  48.  Histograms  of  the  distribution  of  the  head  length  expressed  as  a  percentage  of  the 
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FIG.  49.  Histograms  of  the  distribution  of  the  pectoral  fin  length  expressed  as  a  percentage 
of  the  standard  length  in  populations  from  various  localities.  Locality  coding  as  in 
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A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


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FIG.  50.     Histograms  of  the  distribution  of  the  pectoral  fin  lengths  for  the  entire  sample 
of  B.  intermedius.     Lengths  are  expressed  as  a  percentage  of  the  standard  length. 


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FIG.  51.     Histogram  of  the  distribution  of  the  lateral  line  count  for  the  entire 
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FIG.  52.     Histograms  of  the  distribution  of  the  lateral  line  scale  counts  of  Barbus  intermedius 
populations  from  various  localities.     Locality  coding  as  in  Fig.  34. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


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FIG.   53.     Histogram  of  the  distribution  of  the  dorsal  spine  lengths,   expressed  as  a 
percentage  of  the  standard  length  of  the  entire  Barbus  intermedius  sample. 


the  longest  heads  (Barbus  brunellii,  fig.  46,  and  Barbus  gorguarii,  fig.  39)  are  usually 
large  mouthed  but  the  correlation  is  not  perfect.  The  head  length  distribution  is 
normal  (fig.  47)  when  the  whole  sample  is  considered,  but  the  small  samples  from 
some  localities,  e.g.  the  Blue  Nile,  give  a  chance  bimodal  or  discontinuous  distri- 
bution (fig.  48).  No  significance  can  be  attached  to  this. 

The  pectoral  fin  is  modally  longest  in  the  Lake  Baringo  sample  (fig.  48).  The 
histogram  for  the  entire  sample  (fig.  50)  is  bimodal,  suggesting  a  significant  difference 
in  the  Baringo  fishes  and  these  (see  below)  have  been  accorded  sub-specific  status. 

Squamation.  The  scales  bear  numerous  longitudinal  striae.  The  lateral  line 
scale  count  range  for  the  whole  sample  is  from  26  to  35  (fig.  51).  The  range  and 
frequencies  for  each  population  show  no  significant  variation  (fig.  52).  The  lateral 
line  count  is  adequate  for  distinguishing  a  Barbus  intermedius  sample  from  a  Barbus 
altianalis  sample,  but  is  of  no  value  for  characterizing  the  subspecies  of  Barbus 
intermedius.  There  are  almost  always  twelve  scales  around  the  caudal  peduncle. 
The  only  exceptions  are  three  specimens  from  Lake  Tsana  that  have  13.  Between 
the  dorsal  mid-line  and  the  lateral  line  there  are  4-5  (f.24),  5  (f.i),  5-5  (f.38i), 
6  (f.2),  or  6-5  (f.33)  scale  rows.  Between  the  lateral  line  and  the  ventral  mid-line 
there  are  4  (f.i),  4-5  (f.i6o),  5  (f.2),  5-5  (1.231),  6  (f.i),  6-5  (f.3o)  or  7-5  (f.i)  scale 
rows.  Between  the  lateral  line  and  the  base  of  the  ventral  fin  there  are  1-5  (f.2), 
2  (1.9),  2-5  (1246),  3  (f.4i),  3-5  (f.i37),  4  (f.i)  or  4-5  (f.3)  scale  rows.  The  number  of 
scale  rows  is,  not  infrequently,  different  on  either  side  of  the  specimen  and  in  many 
cases  scales  had  been  lost  and  it  was  impossible  to  count  the  original  number. 

Zolezzi  (1940)  gives  39-40  scales  in  the  lateral  line  series  of  Barbus  platystomus 
var.  vatovae.  The  only  specimen  in  the  collections  of  the  Stazione  Idrobiologia, 
Roma  (where  Zolezzi's  specimens  were  lodged),  that  is  attributed  to  this  variety  and 
is  of  the  same  length  as  Zolezzi's  holotype  has  only  30  lateral  line  scales  on  one  side 
and  31  on  the  other.  This  specimen  is  not  labelled  as  type  material  but  agrees  with 
the  only  specimen  described  by  Zolezzi  in  all  respects  except  for  the  lateral  line  scale 
count. 

Dorsal  fin.  The  origin  of  the  dorsal  fin  varies  from  just  in  front  to  just  behind 
the  vertical  to  the  origin  of  the  pelvic  fins. 

There  are  four  unbranched  rays.  The  last  of  these  is  ossified  into  a  smooth  spine, 
x  =  20-1,  s.d.  =  4-9,  s.e.  =  0-2,  range  6-9-33-9  (percentage  of  the  S.L.).  The 
range  is  unusually  large  and  reflects  the  size  range  of  the  specimens  examined  and 


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FIG.  56.  A  comparison  of  the  left  pharyngeal  bones  from  a  shallow-bodied  specimen  of,  A, 
B.  intermedius  (previously  identified  as  B.  gorguarii)  of  329  mm  S.L.  with  a  deep-bodied 
specimen,  B,  (previously  identified  as  B.  macronema)  of  332  mm  S.L. 


the  negative  allometry  of  the  dorsal  fin  spine  (see  below).  A  histogram  of  the  dis- 
tribution of  the  length  of  the  dorsal  fin  spine  (fig.  53)  shows  that  the  whole  sample  is 
bimodal.  The  fishes  with  the  relatively  longer  spines  all  came  from  Lake  Baringo 
(fig.  54).  The  length  of  the  spine  serves  as  one  of  the  distinguishing  characters  of 
this  population  (see  above).  "Without  the  Lake  Baringo  fishes  the  dorsal  spine  has 
a  mean  length  of  19-1  and  the  same  range  as  in  the  whole  sample  (6-9-33-9).  These 
values  cover  too  wide  a  range  to  be  useful  as  a  diagnostic  character.  A  graph  of 
the  dorsal  spine  length  (expressed  as  a  percentage  of  the  S.L.)  against  the  standard 
length  (fig.  55)  shows  that  the  spines  are  relatively  shorter  in  longer  fishes.  For  fish 
of  less  than  170  mm  S.L.  the  dorsal  spine  values  are  x  =  22-6;  s.d.  =  3-4;  s.e.  =  0-3; 
range  16-3-33-7  '>  f°r  fishes  of  171-250  mm  S.L.,  x  =  18-9;  s.d.  =  2-3;  s.e.  =  0-2; 
range  13-8-24-5;  for  fishes  of  more  than  251  mm  S.L.,  x  =  14-2;  s.d.  =  2-4; 
s.e.  =  0-2;  range  =  6-9-19-7. 

Almost  the  same  number  of  specimens  have  eight  branched  dorsal  fin  rays  as 
have  nine.  No  other  number  was  observed  except  in  cases  which  were  obviously 
the  result  of  physical  damage. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


69 


FIG.  57.     The  pharyngeal  bones  from  a  specimen  of  Barbus  intermedius 
(previously  identified  as  B.  surkis)  of  348  mm  S.L. 


The  anal  fin  invariably  has  three  unbranched  rays  and  five  branched  rays. 

The  gill  raker  count  was  only  taken  on  47  specimens.  The  range  is  from  10  to  16 
on  the  lower  arm  of  the  first  gill  arch.  The  only  factor  which  might  have  some 
significance  is  that  the  long-headed  fishes  tend  to  have  10-12  gill  rakers  whilst  the 
shorter-headed  fishes  have  12-16  gill  rakers.  Not  enough  specimens  were  examined 
to  attribute  any  real  significance  to  this  difference. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  bones  vary  in  shape.  Generally  at 
an  equal  S.L.  fishes  with  a  proportionately  longer  head  have  thinner  pharyngeal 
bones  than  shorter-headed  specimens  ;  fig.  56  contrasts  the  pharyngeals  of  a  speci- 
men of  the  ' gorguarii'  form,  a  fish  of  329  mm  S.L.,  with  the  pharyngeals  of  a  specimen 
of  the  'macronema'  form  of  323  mm  S.L. 

An  even  greater  difference  is  immediately  noticeable  in  the  deep-bodied  speci- 
mens ;  fig.  57  shows  the  pharyngeals  from  a  specimen  of  the  '  surkis '  form  of 
348  mm  S.L.  The  bone  in  this  case  is  extremely  thick  and  the  teeth  are  molariform. 
A  radiograph  of  this  specimen  [B.M.  (N.H.)  No.  1902.12.13  :  229]  shows  its  stomach 
to  be  full  of  gastropod  mollusc  shells.  Possibly  the  difference  in  pharyngeal  bone 
strength  reflects  differences  in  diet  and  age.  The  left  pharyngeal  bone  of  a  slender- 
bodied  Barbus  brunellii  is  shown  in  fig.  58. 

The  pharyngeal  teeth  number,  without  exception,  2. 3.5. -5.3. 2.  In  fishes 
<  180-200  mm  S.L.,  the  teeth  are  long  with  recurved,  mammilliform  crowns  (as 
shown  in  fig.  58)  whilst  larger  and  bulkier  fishes  have  teeth  like  those  shown  in  fig. 
57.  Needless  to  say  these  examples  are  linked  by  an  almost  continuous  series  of 
intermediate  shapes.  I  am  unable  to  offer  anything  more  than  the  most  tentative 
elucidation  of  this  variation  (see  p.  127). 


K.    E.    BANISTER 


FIG.  58.     Pharyngeal  bones  from  a  specimen  of  Barbus  intermedius 
(one  of  Bini's  B.  brunellii). 

Little  is  known  about  the  effect  of  age  and  diet  upon  cyprinid  pharyngeal  teeth. 
The  range  in  pharyngeal  tooth  form  shown  by  Barbus  intermedius  is  paralleled  to 
some  extent  by  that  in  Barbus  altianalis  where  the  deep-bodied  form  hitherto 
referred  to  Barbus  obesus  possesses  singularly  massive  pharyngeal  bones  (see  p.  14). 

Coloration.  This  is  very  variable.  In  life  the  body  colour  ranges  from  silvery- 
grey  to  blue,  through  green,  olive  and  brown,  to  bronze.  Some  specimens  have 
been  reported  with  pinkish  or  yellowish  bellies.  The  fins  have  been  described  as 
whitish  tinged  with  crimson,  deep-green,  slaty-grey,  yellowish  and  yellow-green.  It 
is  quite  possible  that  the  colour  may  be  a  constant  difference  distinguishing  popu- 
lations, but  there  is  not  enough  information  available  at  the  moment.  In  alcohol- 
preserved  specimens  the  body  is  grey,  brown  or  bronzy,  the  scales  are  usually  darker 
at  the  base  and  the  fins  more  or  less  the  colour  of  the  body. 

DISTRIBUTION.  Barbus  intermedius  is  widely  distributed  throughout  Southern 
Ethiopia  and  into  Northern  Kenya,  certainly  as  far  as  Lake  Baringo  (fig.  60).  It  has 
been  recorded  from  the  following  rivers  :  Omo  system  -  Zendo,  Gibe,  Wondinak 
and  Ergino  rivers  ;  Webi  Shebeli  system  -  Errer,  Jerrer,  Iraro  and  Modjo  rivers  ; 
Hawash  system  -  Kassam,  Hurso,  Akaki  and  Gota  rivers  ;  Blue  Nile  system  - 
Wama,  Urgessa,  Gudar,  Metti,  Juju,  Didessa  and  Mogre  rivers  ;  Euasso  Nyiro 
system  -  Uaso  Narok  ;  Juba  system  -  Awata  river  ;  Lake  Zwai  basin  -  Suksuki 
and  Maki  rivers  ;  Lakes  Abaya  and  Ganjule  basins  -  Zeissi,  Sire,  Ganda,  Elgo, 
Alaba  and  Burka  rivers  ;  Lake  Stephanie  basin  -  Zuja,  Sagan,  Gato  and  Bar] a 
rivers  ;  Lake  Rudolf  basin  -  Kerio  and  Ngeng  rivers.  It  has  also  been  recorded 
from  Lakes  Zwai,  Tsana,  Baringo,  Langano,  Abaya,  Ganjule,  Orsodi  and  Stephanie. 

There  has  been  a  certain  amount  of  confusion  over  the  presence  of  Barbus  gregorii 
(=  Barbus  intermedius  australis)  in  the  Athi-Tana  system  as  Mann  (1971)  noted. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  71 

The  localities  for  the  syntypes  are  given  by  Boulenger  (191  la)  as  :  i  Kiroruma 
(Upper  Tana),  Leikipia  ;  2-4,  Guasso  el  Narua  (Baringo),  Leikipia  ;  5-6,  Guasso 
Nyuki  (Naiwasha),  Njemps  Ndogo  and  a  skeleton,  7,  Kibwesi  river  (Athi).  All  of 
these  specimens  were  collected  by  Professor  J.  W.  Gregory's  expedition.  Mann 
(ibid.)  was  unable  to  trace  these  localities  with  any  certainty,  but  he  points  out  that 
the  Leikipia  plateau  drains  into  the  northern  Euasso  Nyiro  and  Lake  Baringo,  not 
into  the  Tana  system.  Gregory  (1896)  in  his  account  of  his  travels  gives  grid 
references  for  Guasso  Nyuki  and  Guasso  el  Narua.  The  former  is  a  small  stream 
at  o°28'  N,  36°o8'  E,  slightly  east  of  south  of  Lake  Baringo,  the  latter  is  at  the  foot 
of  the  Leikipia  escarpment,  to  the  east  of  Lake  Hannington  at  o°i6'  N,  38°i8'  E. 
Guasso  Nyuki  is  nowhere  near  Lake  Naivasha.  The  map  of  Gregory's  route  crosses 
these  rivers  at  the  grid  references  given.  The  problem  of  Kiroruma  is  not  so  easily 


FIG.  59.     The  distribution  of  B.  intermedius  :    A  =  B.  intermedius  intermedius, 
•  =  B.  intermedius  australis. 


72  K.    E.  BANISTER 

settled.  Mann  (ibid.)  says  that  it  is  not  on  any  modern  maps.  Gregory's  (loc.  cit.} 
gazetteer  gives  the  following  information  ;  Kiroruma  =  Kiloluma  o°4o'  S,  37°3o'  E. 
The  river  is  marked  on  his  map  and  is  in  the  upper  Tana  system,  parallel  to  the 
Thika-Thika  river,  Gregory's  route  did  not  pass  through  that  grid  reference  (accord- 
ing to  his  route  map)  and  at  the  nearest  was  24  km  away  from  his  location  of  the 
Kiroruma  river.  From  Gregory's  text  (1896  :  199)  it  appears  that  he  was  in  a  great 
hurry  at  the  time  that  he  was  in  this  area  and  there  is  no  mention  of  specimens 
having  been  collected.  The  Kiroruma  is  separated  from  the  Leikipia  escarpment 
and  plateau  by  Mount  Kenya  and  no  specimens  of  Barbus  intermedius  (sensu  lato) 
have  been  collected  from  neighbouring  rivers  in  the  Tana  system  (but  see  below). 
At  the  moment  this  matter  cannot  be  resolved. 

Specimen  No.  7  of  Barbus  gregorii  in  Boulenger  (191  la  :  46)  from  Kibwesi  is 
another  problem.  It  bears  the  B.M.  (N.H.)  Reg.  No.  1893.12.2  :  36.  This  number, 
according  to  the  register,  is  one  of  16  specimens  of  Barbus  tanensis  (  =  Barbus 
oxyrhynchus)  brought  back  by  the  Gregory  expedition.  Barbus  intermedius  and 
Barbus  tanensis  were  the  only  Barbus  spp.  brought  back  by  the  expedition  and  they 
are  easy  to  distinguish,  especially  so  as  the  specimens  are  large.  The  other  five 
specimens  bearing  the  locality  Kibwesi  are  undoubtedly  Barbus  tanensis.  This  is 
inexplicable.  Three  of  Gregory's  specimens  of  Barbus  tanensis  were  recorded  as 
having  come  from  Guasso  el  Narua  (see  above).  This  species  has  not  been  found 
there  since  and  I  am  at  a  loss  to  explain  its  alleged  presence  there.  Perhaps  the 
locality  was  incorrectly  recorded,  there  are  enough  inconsistencies  in  this  collection 
to  cast  a  shadow  of  doubt  over  some  of  the  localities.  The  Lake  Baringo  basin  is 
the  most  southerly  definitive  record  of  Barbus  intermedius,  but  further  south  in  the 
rift  valley  before  the  Aberdare  mountains  and  the  Maui  (or  Mau)  escarpment  lie 
Lakes  Hannington,  Elementaita,  Nakuru  and  Naivasha.  The  first  three  of  these 
are  extremely  alkaline.  Lake  Naivasha  according  to  Worthington  (i932b,  1933) 
and  Copley  (1948)  has  a  small,  cyprinodont,  Aplocheilichthys  antinorii,  as  its  only 
indigenous  fish,  but  Tilapia  nigra  was  introduced  in  1925  and  Micropterus  salmoides 
was  introduced  later.  Copley  (1948)  mentions  that  no  Barbus  species  are  present 
in  Lake  Naivasha  but  Parenzan  (1939)  lists  Barbus  gregorii  ( =  Barbus  intermedius) 
as  present  in  the  lake.  Whether  this  locality  of  Parenzan's  was  based  on  first-hand 
evidence  or  taken  from  Boulenger's  (igna)  list  I  cannot  find  out. 

The  significance  of  Aplocheilichthys  antinorii  is  that  it  also  lives  in  Southern 
Ethiopia,  i.e.  the  distribution  is  similar  to  that  of  Barbus  intermedius.  Cooke  (1958), 
in  his  reconstructions  of  the  lower  Pliocene  drainage  of  east  Africa,  shows  volcanic 
highlands  present  in  the  Aberdare  region  (to  the  immediate  south  of  Lake  Naivasha) 
and  the  'Nile'  and  'Athi-Tana'  drainage  areas  much  as  they  are  today.  This 
ancient  separation  ol  the  two  watersheds  largely  precludes  the  presence  of  many 
species  in  common. 

The  northern  Euasso  Nyiro  river  presents  certain  problems,  not  the  least  of  which 
is  the  paucity  of  specimens.  From  this  river  in  the  region  of  Chanlers  Falls  come 
the  three  Barbus  erlangeri  ( =  Barbus  intermedius)  specimens  whilst  from  the  eastern 
extremity  of  the  system  in  the  Nero-Narok  and  Ngau-Narok  systems  associated 
with  the  Lorian  swamp  come  17  specimens  of  Barbus  oxyrhynchus.  The  presence  of 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS 
90- 


73 


80- 

70- 

1  ,      "     ' 

60- 
50- 

i 

40- 

30-              -TTTTTTTT 

20- 

i 

10- 

i 

1  '           1 

I'NIr!      i     1 

I               i  i                 !  !  '  :     | 

26      27      28      29      30      31       32      33      34      35      36 


r 


10- 
20- 
30- 
40- 
50- 

60- 

FIG.  60.     Histograms  comparing  the  distribution  of  the  lateral  line  counts  of 
Barbus  intermedius  (above)  with  B.  altianalis  (below). 

'  Nilotic '  and  an  '  Athi-Tanan '  species  in  the  same  river  system  is  surprising,  but  the 
watersheds  across  the  divide  are  low.  This  area  is  now  semi-desert,  but  about  the 
time  that  Lake  Baringo  was  connected  to  Lake  Rudolf  the  water  table  must  have 
been  much  higher  and  water  connections  must  have  linked  the  two  river  systems  in 
this  region. 

The  subsequent  drop  of  water  level  has  left  the  Euasso  Nyiro  system  as  a  relict 
area  containing  examples  of  the  formerly  contiguous  faunae. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  intermedius  is  a  variable  species  and  bears  a 
close  gross  morphological  resemblance  to  Barbus  altianalis,  to  which  I  suggest  it  is 
very  closely  related.  These  two  species  (and  Barbus  acuticeps  and  Barbus  ruasae] 
form  a  group  of  closely  related  species  referred  to  here  as  the  intermedius  group  or 
complex. 

Typically  Barbus  intermedius  has  a  shallow  compressed  body,  a  caudal  peduncle 
longer  than  it  is  deep,  numerous  more  or  less  parallel  striae  on  the  scales  and  a  well- 
ossified  dorsal  spine  of  moderate  length  and  stoutness. 


74  K.    E.    BANISTER 

It  is  difficult  to  distinguish  a  specimen  of  Barbus  altianalis  from  Barbus  intermedium 
if  the  locality  is  unknown.  There  are  modal  differences  in  the  populations  which 
are  listed  under  Barbus  altianalis. 

Both  species  are  polytypic  and  can  show  considerable  variation,  particularly  in 
body  shape  and  mouth  form.  However,  it  has  always  proved  possible  to  distinguish 
an  unusually  deep-bodied  member  of  the  Barbus  intermedius  group  from  a  member 
of  the  typically  deep-bodied  Barbus  bynni  group  because  members  of  the  latter 
group  have  more  compressed  bodies  and  longer,  stronger  dorsal  spines. 

Barbus  intermedium  is  not,  as  far  as  I  know,  found  in  the  same  localities  as  Barbus 
altianalis.  The  nearest  approach  of  these  two  species  appears  to  be  in  the  poorly 
collected  Karasuk  area  to  the  north-east  of  Lakes  Victoria  and  Kioga  where  the 
headwaters  of  the  Lake  Rudolf  basin  streams  and  the  Lake  Victoria  and  Kioga 
basin  streams  are  very  close. 

The  intermedium  complex  is  discussed  further  on  p.  128.  The  two  subspecies  of 
Barbus  intermedium  can  be  distinguished,  not  only  by  locality,  but  also  by  the  modally 
much  longer  dorsal  spine  of  Barbum  intermedium  australis  (x  =  24-9  against  19-1  in 
Barbus  intermedium  intermedium}.  Barbus  intermedium  australis  also  has  a  longer 
pectoral  fin  (x  =  23-4  against  21-5)  and  longer  barbels  (Ab,  x  =  7-1  against  5-6  ; 
Pb,  x  =  8-4  against  6-7).  Regrettably  it  is  not  always  possible  to  place  an  individual 
specimen,  lacking  locality  data,  in  the  correct  group. 


Barbus  intermedius  intermedius  Riippell 

A  general  description  is  given  on  p.  51.  Them  orphometric  data  for  this  sub- 
species are  : 

n  t    x  s.d.  s.e.  range 

D  388  28-1  2-8  o-i  20-7-37-7 

H  388  26-5  2-3  o-i  19-0-31-7 

I  388  6-0  1-4  o-i  3-5-10-2 

IO  388  8-0  0-9  0-04  4-7-11-6 

MW  386  5-9  i-o  0-05  3-6-  9-1 

Pet  388  21-5  2-0  o-i  15-6-29-0 

CP1  388  17-0  1-6  o-i  11-7-22-9 

CPd  388  11-5  1-2  o-i  8-6-15-0 

Snt  388  8-6  1-2  0-05  5-4-12-7 

Ab  385  5-6  1-8  o-i  i-8-ii-i 

Pb  387  6-7  2-1  o-i  1-9-12-1 

DSp  385  19-1  4-2  0-2  6-9-33-9 

Not  all  specimens  examined  are  included  in  the  morphometric  data  above. 

The  size  range  of  the  specimens  is  94  to  489  mm  S.L.  The  lateral  line  count 
ranges  from  26  to  34;  26  (f.7),  27  (f.27),  28  (f.49),  29  (f.85),  30  (1.85),  31  (£63), 
32  (f.36),  33  (f.25),  34  (f.5). 

Between  the  dorsal  mid-line  and  the  lateral  line  there  are  4-5  (f.24),  5  (f.i), 
5'5  (f-325)>  6  (f.2)  or  6-5  (f.32)  scale  rows.  Between  the  lateral  line  and  the 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  75 

ventral  mid-line  there  are  4  (f.i),  4-5  (f.iog),  5  (f.2),  5-5  (f.226),  6  (f.i),  6-5  (£.29)  or 
7-5  (f.i)  scale  rows.  Between  the  lateral  line  and  the  base  of  the  ventral  fin  there 
are  1-5  (f.2),  2  (£.9),  2-5  (£.242),  3  (£.40),  3-5  (£.85),  4  (f.i)  or  4-5  (£.3)  scale  rows. 

DISTRIBUTION.     The  distribution  is  as  on  p.  70  except  for  Lake  Baringo. 


Barbus  intermedius  australis  ssp.  nov. 

HOLOTYPE.  A  fish  of  128  mm  S.L.,  No.  18  in  1932.6.13  :  191-200,  from  a  jar 
labelled  Barbus  gregorii,  Lake  Baringo,  in  the  collection  of  the  B.M.  (N.H.).  This 
specimen  was  selected  as  being  close  to  the  mean  for  most  morphometric  characters, 
and  therefore  is  typical  of  the  population. 

DESCRIPTION.  The  description  is  largely  as  for  the  nominate  subspecies  (see 
p.  51).  The  morphometric  data  in  detail  are  as  follows  and  based  on  58  specimens, 
S.L.  66-388  mm. 

n  x  s.d.  s.e.  range 

L  66    -388  mm 

D  58  30.0  2-2  0-3  26-3-35-1 

H  58  26-2  1-7  0-2  23-0-31-0 

I  58  6-0  i-o  o-i                  4-4-  9-1 

IO  58  8-1  i-o  o-i                  6-1-12-3 

MW  58  5-5  0-7  o-i                  4-1-  7-3 

Pet  58  23-4  1-5  O-2  2I-O-27-2 

CP1  58  16-1  1-4  0-2  11-7-18-7 

CPd  58  12-7  0-9  o-i  9-5-14-2 

Snt  58  8-4  0-9  o-i  6-7-10-7 

Ab  58  7-1  1-2  0-2  4-8-10-0 

Pb  58  8-4  1-3  0-2  6-0-10-9 

DSp  58  24-9  3-9  0-5  15-6-32-3 

The  number  of  scales  in  the  lateral  line  ranges  from  27  to  33  :  27  (£.4),  28  (f.n), 
29  (f.i8),  30  (£.9),  31  (f.8),  32  (f.6),  33  (f.2).  Only  one  specimen  has  6-5  scale  rows 
between  the  dorsal  mid-line  and  the  lateral  line,  the  rest  have  5-5.  Between  the 
lateral  line  and  the  ventral  mid-line  there  are  4-5  (£.51),  5-5  (£.4),  or  6-5  (f.i)  scale 
rows.  Between  the  lateral  line  and  the  base  of  the  pelvic  fin  there  are  2-5  (f-4), 
3  (f.i),  or  3-5  (£.52)  scale  rows. 

The  distinguishing  characters  of  the  two  subspecies  are  the  longer  dorsal  spine, 
longer  pectoral  fins,  slightly  deeper  body  and  longer  barbels  in  Barbus  intermedius 
australis. 

Lake  Baringo  is  an  alkaline  lake,  and  it  is  impossible  to  say  whether  the  Baringo 
population  displays  its  particular  phenotype  as  a  result  of  the  environment  or  as  a 
result  of  the  genotype.  There  are  very  slight  indications  that  the  fishes  from  the 
Omo  river  and  Lake  Rudolf  incline  slightly  towards  the  Baringo  facies  but  it  must 
be  remembered  that  the  Lake  Rudolf  and  Omo  river  sample  is  extremely  small. 

DISTRIBUTION.     This  species  is  known  from  Lake  Baringo,  Kenya. 


76  K.   E.    BANISTER 

Barbus  longifilis  Pellegrin  1935 

Barbus  altianalis  var.  longifilis  Pellegrin,  1935,  Revue  Zool.  Bot.  afr.  27  (3)  :  376-385  (part,  not 
the  specimen  from  Nyabarongo). 

NOTES  ON  THE  DETERMINATION.  This  species,  from  the  upper  reaches  of  the 
Luhoho  Congo,  is  not  a  variety  of  Barbus  altianalis,  but  a  well-defined  species.  One 
of  Pellegrin's  type  series  (M.H.N.P.  No.  35-75)  is  better  referred  to  Barbus  pauci- 
squamatus.  Poll  (1939 : 69)  synonymized  Barbus  altianalis  var.  longifilis  with 
Barbus  altianalis  var.  paucisquamata,  a  move  which  does  not  take  into  account  the 
much  longer  dorsal  spine,  the  much  longer  barbels  and  the  much  deeper  body  of  the 
former  variety. 

LECTOTYPE.  A  fish  of  173  mm  S.L.  (M.H.N.P.  No.  35-150).  This  is  the  least- 
damaged  specimen  of  Pellegrin's  type  series  from  the  Loama  river. 

DESCRIPTION.     The  description  is  based  on  nine  specimens,  132-247  mm  S.L. 

x  s.d.  s.e.  range 

D  31-2  1-6  0-5  29-2-34-0 

H  24-6  0-7  0-2  23-5-25-3 

I  6-5  0-5  o-i  6-0-  7-2 

IO  8-0  0-5  0-2  7-0-  8-6 

MW  5-5  0-6  0-2  4-5-  6-3 

Pet  23-1  i-o  0-3  21-7-24-2 

CP1  17-0  i-o  0-3  15-8-18-3 

CPd  11-4  0-7  0-2  9-9-12-1 

Snt  8-3  0-6  0-2  7-5-  9-3 

Ab  8-9  0-7  0-2  8-3-  9-6 

Pb  11-3  i-i  0-4  9-5-13-4 

Barbus  longifilis  has  a  pointed  snout.  Except  for  a  slight  nuchal  hump  the  dorsal 
profile  is  straight  from  the  snout  to  the  origin  of  the  dorsal  fin.  The  mouth  is  ventral 
with  thin  fleshy  lips  ;  the  barbels  are  characteristically  long. 

Dorsal  Jin.  Has  IV-g  (f.8)  or  IV- 10  (f.i)  rays,  the  last  unbranched  ray  is  solidly 
ossified  into  a  thick  straight  spine  (x  =  25-6,  s.d.  =  2-7,  s.e.  =  i-o,  range  22-0- 
30-3).  The  dorsal  fin  origin  is  usually  in  front  of  the  insertion  of  the  pelvic  fins.  A 
low  sheath  of  large  scales  surrounds  the  base  of  the  dorsal  fin. 

The  anal  fin  has  three  unbranched  and  five  branched  rays. 

Squamation.  The  striae  on  the  scales  are  slightly  irregular,  either  parallel  or 
slightly  converging,  the  lateral  line  has  25  (f.i),  26  (f.3),  27  (£3)  or  28  (f.2)  scales. 
There  are  4-5  scales  between  the  dorsal  mid-line  and  the  lateral  line  and  5-5  between 
the  lateral  line  and  the  ventral  mid-line.  There  are  2-5  (f-7)  or  3  (f.2)  scale  rows  be- 
tween the  lateral  line  and  the  base  of  the  pelvic  fins  and  without  exception  12  scales 
are  present  around  the  caudal  peduncle. 

There  are  between  10  and  12  gill  rakers  on  the  lower  arm  of  the  first  gill  arch  in 
the  specimens  examined. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  tooth  formula  is  2. 3. 5. -5. 3. 2.  The 
first  tooth  in  the  inner  row  (fig.  62)  is  conical  and  directed  posteriorly,  the  second 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


77 


s 


K.   E.    BANISTER 


FIG.  62.     The  left  pharyngeal  bone  from  the  lectotype  of  Barbus  longifilis. 

tooth  is  taller  and  more  flattened  laterally  than  the  first.  A  small  concavity  on  the 
posterior  face  of  the  crown  creates  a  slight  hook,  the  hook  and  the  concavity  pro- 
gressively enlarge  on  teeth  3,  4  and  5.  The  teeth  of  the  second  and  third  rows  are 
small,  less  extreme  versions  of  the  posterior  tooth  on  the  inner  row. 

Coloration.  The  body  of  preserved  specimens  is  brassy,  slightly  darker  on  the 
back  than  on  the  belly.  A  band  of  dark-brown  pigment  is  present  on  the  middle 
third  of  the  fin  membrane  of  the  dorsal  and  pelvic  fins  and  extends  from  the  middle 
to  the  end  of  the  pectoral,  anal  and  caudal  fins,  the  density  of  the  pigment  varies 
considerably  from  specimen  to  specimen,  especially  the  caudal  fin. 

DISTRIBUTION.  The  specimens  examined  came  from  the  Loama  and  Kansehete 
rivers,  Congo. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  longifilis  is  a  member  of  the  Barbus  bynni 
group  (see  p.  36).  The  much  longer  barbels  distinguish  this  species  from  Barbus 
bynni  and  Barbus  gananensis.  In  other  respects  it  most  closely  resembles  Barbus 
oxyrhynchus  and  were  the  two  species  found  in  the  same  area  it  may  be  difficult  to 
assign  some  individual  specimens  (especially  preserved  material)  to  either  of  the  two 
species.  The  longer  barbels  and  more  heavily  pigmented  fins  in  Barbus  longifilis 
will  usually  enable  it  to  be  distinguished  from  Barbus  oxyrhynchus  where  locality 
data  are  unavailable. 

Cooke  (1958  :  26)  presents  some  evidence  to  suggest  that  some  Athi-Tana  faunal 
elements  had  been  able  to  move  across  the  Lake  Victoria  area.  He  cites  the  presence 
of  the  Athi  river  species  Tilapia  nigra  which  occurs  in  the  mid-Pleistocene  beds  at 
Rawi  in  the  Kavirondo  Gulf.  The  relationship  of  Lake  Victoria  to  the  headwaters 
of  the  Congo  in  Pleistocene  times  is  discussed  on  page  22.  There  is  just  a  possibility 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS 


79 


that  there  could  have  been  a  movement  of  Barbus  oxyrhynchus  (or  its  ancestor)  from 
the  Athi  into  the  Congo  before  the  rift  valley  broke  the  connection  (see  fig.  4  in 
Cooke,  op.  cit.)  and  that  Barbus  longifilis  represents  a  surviving  population  descended 
from  this  migration. 

Barbus  macrolepis  Pfeffer  1889 

Barbus  macrolepis  Pfeffer,  1889,  Jb.  hamb.  wiss.  Anst.  6  (2)  :  17  ;    Pfeffer,  1893,  Jb.  hamb.  wiss. 
Anst.  10  (2)  :  34,  pi.  i,  fig.  i  ;    Pfeffer,  1896,  Thierw.  O-Afr.  Fische  :  63. 

LECTOTYPE.  The  lectotype  was  selected  by  Ladiges  et  alii  (1958)  as  a  fish  of 
108  mm  S.L.  (this  specimen  was  kindly  measured  for  me  by  Dr  W.  Ladiges),  Reg. 
No.  H.330  from  Mbusini  on  the  Wami  River,  Tanzania. 

DESCRIPTION.     The  description  is  based  on  20  fishes,  from  48  to  243  mm  S.L. 


L 
D 
H 

I 

10 

MW 

Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


x 


27-6 

7-6 

8-9 

6-8 

21-7 

14-6 

14-8 

7-9 

3'5 
5-8 


s.d. 

3'5 
2-3 
i-4 
0-9 
0-8 
1-6 
1-9 

i'4 

1-7 

i'3 
2-5 


s.e. 

0-8 
o-5 
o-3 

0-2 
0-2 

0-4 
0-4 

o-3 
0-4 

o-3 
0-6 


range 

48    -243  mm 

29-9-38-2 

24-4-31-3 

4-6-10-1 

7.3-10-4 

5-3-  8-3 
18-5-24-6 
11-4-20-3 
12-9-17-2 

5-8-  9-0 

1-6-  6-3 

1-4-  9-0 


FIG.  63.     Barbus  macrolepis  (from  Boulenger  igua). 


8o 


K.   E.   BANISTER 


5  mm 


FIG.  64.     A  scale  from  the  shoulder  of  B.  macrolepis  to  show  the  position  of  the  striae. 


The  three  paralectotypes  examined  have  the  overall  body  shape  of  the  figured 
example.  The  larger  fish  have  a  straighter  ventral  profile  and  a  more  humped 
back.  The  mouth  is  sub-terminal  or  just  ventral,  horse-shoe  shaped  ;  in  all  the 
specimens  the  lips  are  slightly  thickened  and  fleshy. 

Of  four  specimens  radiographed,  three  had  36  vertebrae  and  one  had  37. 

Dorsal  fin.  Has  four  unbranched  rays  and  9  (f.y)  or  10  (f.i3)  branched  rays. 
The  last  unbranched  ray  is  weakly  ossified  with  persistent  articulations  distally. 
The  length  of  the  non-articulated  part  ranges  from  8-7  to  20-8  per  cent  S.L.  with  a 
mean  value  of  15-3.  The  whole  ray  though,  when  unbroken,  is  35  per  cent  of  the 
S.L.  which  gives  a  high  dorsal  fin,  with  a  markedly  concave  dorsal  margin.  This 
ray  is  more  elongated  in  larger  fish.  The  syn types  have  a  very  small  sheath  of  scales 
at  the  base  of  the  dorsal  fin.  -This  sheath  is  not  present  in  the  larger  fish  ;  as  there 
is  no  sign  of  physical  damage  it  presumably  has  been  lost  as  a  result  of  growth.  The 
leading  edge  of  the  dorsal  fin  is  slightly  in  advance  of  the  pelvic  fin. 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays.  The  last  ray  of 
the  latter  is  in  some  cases  markedly  bifurcated. 

Squamation.  There  are  22  (f.7),  23  (f.5),  24  (f.4)  or  25  (f.i)  scales  in  the  lateral 
line  ;  3-5  (f.6)  or  4-5  (f.i2)  scales  between  the  dorsal  mid-line  and  the  lateral  line 
and  3  (f.i),  3-5  (f-7)  or  4-5  (f.i)  scales  between  the  lateral  and  the  ventral  mid-line. 
On  several  specimens  the  scales  could  not  be  counted  reliably.  One  and  a  half  (f.4) 
or  2-5  (f.i6)  scale  rows  are  present  between  the  lateral  line  and  the  insertion  of  the 
pelvic  fin.  There  are  12  scales  around  the  caudal  peduncle. 

The  striae  on  the  scales  are  characteristic  (fig.  64).  They  are  comparatively 
sparse  on  the  exposed  portion  of  the  scale  and  converge  towards  the  posterior  edge 
of  the  scale.  In  this  respect  they  contrast  significantly  with  those  of  Barbus 
oxyrhynchus  (fig.  81). 

Pharyngeal  bones  and  teeth  (figs.  65  and  66).  The  first  tooth  of  the  inner  row  is 
small,  conical  with  a  small  spoon-shaped  depression  at  the  crown.  The  second 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


81 


tooth  is  much  thicker  and  higher  with  a  hooked  crown.  The  teeth  become  pro- 
gressively thinner  posteriorly  and  the  hook  and  the  depression  become  more  exag- 
gerated. The  pharyngeal  tooth  formula  is  2.3. 5. -5.3. 2. 

Gill  rakers.  There  are  12-14  broad,  hooked  gill  rakers  on  the  ventral  limb  of  the 
first  gill  arch. 

Coloration.  Preserved  specimens  are  light  brown  on  the  back  and  a  paler  silvery- 
brown  on  the  lower  part  of  the  flanks.  The  scales  are  dark  edged.  The  caudal 
and  dorsal  fins  are  brownish,  the  other  fins  are  hyaline. 


FIG.  65.     The  left  pharyngeal  bone  from  a  specimen  of  B.  macrolepis  of  94  mm  S.L. 


FIG.  66.     The  left  pharyngeal  bone  from  a  specimen  of  B.  macrolepis  of  243  mm  S.L. 


82 


K.   E.    BANISTER 


Juba 


FIG.  67.     A  map  of  the  distribution  of  Barbus  macrolepis. 


DISTRIBUTION.  This  species  is  known  from  Katare  (or  Kotare),  Malagarasi 
swamp  ;  from  the  Wami,  Ruaha  and  Rufiji  rivers,  Tanzania. 

The  distribution  of  this  species  is  rather  unusual  in  that  it  is  confined  to  a  narrow 
belt  between  5  and  8  degrees  South  but  covers  a  wide  longitudinal  range  from 
Malagarasi  to  near  the  Tanzanian  coast.  The  Malagarasi  river  drains  westwards  into 
Lake  Tanganyika  whilst  the  other  two  rivers  flow  eastward  into  the  Indian  Ocean. 

The  Malagarasi  has  certain  Congo  faunal  affinities  and  its  geological  history 
suggests  that  it  was  once  part  of  the  Congo  system.  The  eastward  flowing  rivers 
do  not  have  any  Congo  links. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  macrolepis  is  a  distinctive  species.  The 
body  is  deep,  the  last  simple  dorsal  fin  ray  is  tall  and  weak,  the  caudal  peduncle  is 
nearly  as  deep  as  it  is  long  and  the  striae  on  the  scales  converge  markedly.  The 
striae  on  the  scales  of  Barbus  marequensis  are  parallel,  but  the  tall  crescentic  dorsal 
fin  of  the  larger  specimens  is  also  present  in  specimens  of  Barbus  marequensis  from 
the  Zambezi  river  to  the  south  of  the  Rufiji  river. 

The  more  heavily  ossified  last  simple  ray  of  the  dorsal  fin  of  Barbus  oxyrhynchus 
in  the  rivers  to  the  north  of  the  Wami  and  the  parallel  sinuous  striae  on  the  scales 
are  easily  distinguished  points  of  difference  from  Barbus  macrolepis. 

Poll  (1967  :  181)  is  of  the  opinion  that  Barbus  jubbi  has  some  marked  similarities 
to  Barbus  macrolepis,  principally  in  the  high  number  of  dorsal  fin  rays  and  the  low 
number  of  scales  in  the  lateral  line  series.  However,  the  striae  on  the  scales  are 
quite  different,  as  is  the  dorsal  fin  spine  and  at  the  moment  I  am  inclined  to  think 
that  the  resemblances  are  spurious. 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  83 

A  very  much  greater  degree  of  morphological  similarity  exists  between  Barbus 
macrolepis  and  Barbus  lagensis  from  Nigeria.  Both  species  have  ten  unbranched 
rays  in  the  dorsal  fin  ;  a  high  but  weak  fourth  unbranched  ray  in  the  dorsal  fin  ;  a 
comparable  low  number  of  scales  in  the  lateral  line  series  and  very  similar  striations 
on  the  scales. 

I  have  not  seen  sufficient  Barbus  lagensis  material  to  comment  further  on  the 
relationships  of  these  two  species  but  both  seem  very  different  from  the  other 
African  Barbus  species  and  if  the  characters  in  common  are  not  the  result  of  con- 
vergence, then  the  real  possibility  exists  that  these  two  species  are  related  and  if  so 
then  their  distribution  may  indicate  that  they  are  relicts  of  an  early  invasion  of 
Barbus  species  (see  p.  132).  I  have  not  seen  any  other  African  Barbus  species  that 
have  converging  striae  on  their  scales  and  the  combination  of  a  deep  body,  large 
scales  and  a  high  dorsal  fin  without  a  well-ossified  dorsal  spine  is  also  unique.  Some 
of  these  characters  are  found  in  some  Asian  Barbus  species,  e.g.  converging  striae 
are  found  in  Barbus  longispinis  Gunther,  this  Celanese  species  has  striae  which 
converge  more  with  age  but  it  has  a  strong  dorsal  spine.  Barbus  macropus  Blkr. 
and  Barbus  huguenini  Blkr.  respectively  from  Borneo  and  Sumatra  are  deep-bodied 
fishes  with  large  scales  and  few  converging  striae  ;  however,  they  have  a  serrated 
dorsal  spine.  I  have  not  been  able  to  find  any  mainland  Asiatic  species  which  have 
all  the  characters  under  discussion,  but  it  seems  that  the  presence  of  converging 
striae  is  more  frequent  in  Asiatic  Barbus  species  than  it  is  in  African  Barbus  species. 
I  have  not  been  able  to  examine  as  many  Indian  species  as  I  would  have  liked  but 
the  occurrence  of  the  converging  striae  in  some  of  the  Asiatic  island  species  suggests 
that  it  may  be  a  primitive  feature  (or  at  least  an  ancient  one)  which  is  present  in 
two  African  species.  At  the  moment  I  cannot  with  any  certainty  align  Barbus 
macrolepis  with  any  Asiatic  species  because  it  is  impossible  to  show  that  the  simi- 
larities in  the  pattern  of  scale  striae  are  not  due  to  convergence.  If  it  is  not  due  to 
convergence,  then  it  is  possible  that  there  may  be  some  fairly  close  relationship 
between  a  group  of  Asiatic  Barbus  species  and  a  pair  of  African  species.  This  idea, 
though,  assumes  that  the  differences  in  the  dorsal  spines  are  of  less  significance  than 
the  similarities  in  the  scale  striations  and  sadly  this  is  a  matter  on  which  I  have  no 
information  at  all. 

Barbus  mariae  Holly  1929 

Barbus  mariae  Holly,   1929,  Anz.  Akad.  Wiss.  Wien  66  (4)  :  34  ;    Copley,  H.,   1958,  Common 

Freshwater  Fishes  of  E.  Africa  :  78-80. 
Barbus  rhinoceros  Copley,  1938,  //.  E.  Africa  Uganda  nat.  Hist.  Soc.,  13  :  191. 

NOTES  ON  THE  SYNONYMY.  It  may  well  eventually  prove  necessary  to  synonymize 
Barbus  mariae  with  Barbus  matris,  Holly,  1928  [Zoo/.  Anz.  Leipzig  85  (1-2)]  from 
the  Athi  river  at  Nairobi.  Holly's  description  of  Barbus  matris  is  very  similar  to 
that  of  Barbus  mariae  but  I  am  unwilling  to  proceed  in  this  matter  without  examin- 
ing the  holotype  of  Barbus  matris  and  at  the  time  of  writing  this  has  not  been 
located. 

LECTOTYPE.  Holly  described  this  species  on  the  basis  of  two  specimens  of 
280  mm  and  295  mm  total  length  from  the  Kitui  river  (Athi  system)  in  Kenya.  I 


84  K.   E.    BANISTER 

have  not  seen  either  of  these  specimens  which  are  supposed  to  be  in  the  Natural 
History  Museum,  Vienna,  but  they  could  not  be  located  by  Dr  P.  Kahsbauer  who 
kindly  searched  for  them.  The  larger  specimen  is  designated  the  lectotype  on  the 
assumption  that  both  specimens  are  extant. 

DESCRIPTION.  The  description  is  based  on  five  specimens  in  the  B.M.  (N.H.) 
collections  of  standard  lengths,  86,  105,  112,  117  and  342  mm  from  the  Athi  river. 

x  range 

D  26-9  24-4-29-5 

H  30-7  30-1-32-5 

I  8-5  5-5-10-4 

10  6-8  5-8-  8-5 

MW  5-2  4-5-  7-0 

Pet  22-5  21-9-22-5 

CP1  17-3  16-1-19-6 

CPd  10-6  9-3-12-9 

Snt  9-9  8-9-10-4 

Ab  2-7  1-2-  3-8 

Pb  5-3  4-5-  5-8 

All  measurements  are  expressed  as  percentages  of  the  standard  length.  With  this 
particular  sample  it  was  not  considered  useful  to  calculate  the  standard  deviation 
and  the  standard  error. 

Barbus  mariae  is  a  distinctive  species.  The  upper  jaw  has  a  remarkable  median 
protrusion  (the  'rhinoceros  horn'  of  Copley  1958)  when  the  mouth  is  open.  This 
is  caused  by  the  fish  having  a  large  kinethmoid  (sensu  Alexander  1966).  The 
antero-ventral  edge  of  the  labial  part  of  the  premaxilla  is  gently  curved  and  over- 
hangs the  lower  jaw  giving  a  'clupeoid'  appearance  to  the  jaws. 

The  body  is  long  and  thin,  the  dorsal  fin  has  its  origin  in  the  posterior  half  of  the 
body  more  or  less  vertically  above  the  insertion  of  the  pelvic  fin. 

Dorsal  fin.  It  has  TV-g  rays.  The  fourth  unbranched  ray  is  heavily  ossified, 
smooth  and  from  17-5  to  32-6,  x  =  29-0,  of  standard  length  (negatively  allometric). 
The  dorsal  margin  of  the  fin  is  markedly  concave. 

Anal  Jin.     With  III-5  rays. 

Squamation.  Holly  gives  30  or  31  for  the  lateral  line  scale  count  of  the  specimens 
he  described.  The  specimens  I  examined  have  27  (f.i),  28  (f.i),  or  29  (f.3)  scales. 
The  difference  is  probably  attributable  to  the  different  techniques  used  in  counting 
these  scales.  The  figures  given  by  Holly  agree  with  the  complete  number  of  scales 
in  the  lateral  line  series  if  27  or  28  are  present  to  the  end  of  the  hypurals.  There 
are  12  scales  around  the  caudal  peduncle,  4-5-5-5  between  the  dorsal  mid-line  and 
the  lateral  line  and  4-5  between  the  lateral  line  and  the  ventral  mid-line.  One  and 
a  half  or  2  scales  were  present  between  the  lateral  line  and  the  base  of  the  pelvic  fin. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  tooth  formula  is  2.3. 5. -5. 3. 2.  The 
first  tooth  in  the  inner  row  is  about  two-thirds  of  the  length  of  the  second.  The 
second  tooth  is  the  widest.  All  in  this  row  are  unicuspid  and  recurved  and  the 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


in 

£ 


cc 

vO 


K.    E.    BANISTER 


FIG.  69.     The  left  pharyngeal  bone  from  a  specimen  of  Barbus  mariae  of  342  mm  S.L. 


teeth  become  more  hooked,  taller  and  thinner  posteriorly.     Only  the  pharyngeal 
teeth  of  the  largest  specimen  were  examined. 

Gill  rakers.  On  the  largest  specimen  9  +  1.  The  gill  rakers  were  widely  spaced 
and  hooked  forward.  This  number  is  lower  than  is  usual  among  the  large  Barbus 
species  under  consideration  in  this  revision. 

Coloration.  Holly's  (op.  cit.)  remarks  on  the  coloration  match  the  colour  of  the 
preserved  specimens  I  examined.  The  back  is  dark  ochre-brown  shading  to  a 
silvery-yellow  in  the  belly.  The  fins  are  very  light  brown  and  the  scales  have  dark 
edges.  Copley  (1958)  states  that  the  live  fish  are  olive-brown  on  the  back  and 
silvery  on  the  belly. 

DISTRIBUTION.  The  five  specimens  1936.12.22  :  35-39  in  the  B.M.  (N.H.)  col- 
lections are  from  the  Athi  river.  Specimen  No.  B.M.  (N.H.)  1937.6.4  :  16  has  only 
the  locality  Kenya.  Copley  (1958)  records  it  from  the  Athi  and  Tana  systems,  he 
also  mentions  that  they  can  weigh  up  to  48  Ib. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  87 

DIAGNOSIS  AND  AFFINITIES.  Barbus  mariae  is  most  easily  distinguished  by  the 
'rhinoceros  horn '-like  development  of  the  kinethmoid  and  the  'clupeoid'  appear- 
ance of  the  upper  jaw.  Barbus  mariae  is  sympatric  with  Barbus  oxyrhynchus  but 
lacks  many  of  its  characters  and  I  am  unable,  at  the  moment,  to  comment  further 
on  its  relationships. 

Barbus  microbarbis  David  and  Poll  1937 

Barbus  microbarbis  David  &  Poll,  1937,  Annls.  Mus.  r.  Congo  Beige  Zool.  (i)  3  (5)  :  261  (only 
the  holotype). 

HOLOTYPE.  A  fish  of  216  mm  S.L.  from  Lake  Luhondo,  Rwanda  ;  M.A.C.T. 
No.  41847.  The  two  paratypes  (M.A.C.T.  Nos.  41848,  41849)  do  not  belong  to 
this  species,  nor  do  any  of  the  other  specimens  in  the  M.A.C.T.  collection  (Nos. 
56449-56455,  56479,  92966-92968,  94219-94220).  This  species  is  split  because 
the  holotype  has  a  sub-terminal  mouth,  five  teeth  in  the  inner  pharyngeal  row,  a 
smoothly  curved  pharyngeal  bone  outline,  nine  branched  rays  in  the  dorsal  fin  and 
15  gill  rakers  on  the  lower  limb  of  the  first  gill  arch  (in  contrast  to  the  ventral  mouth, 
horny  covered  lower  jaw,  four  teeth  on  the  inner  pharyngeal  row,  an  oddly  shaped 
pharyngeal  bone,  10  branched  dorsal  fin  rays  and  19-20  gill  rakers  on  the  lower  limb 
of  the  first  gill  arch  of  the  paratypes  and  the  other  M.A.C.T.  specimens).  All  the 
specimens  previously  included  in  this  species  are  now  considered  to  belong  to 
Varicorhinus  ruandae  Pappenheim,  &  Boulenger,  1914. 

DESCRIPTION.  The  description  is  based  on  the  holotype.  D  =  31-0  ;  H  =  23-6  ; 
I  =  5-6  ;  IO  =  9-2  ;  MW  =  7-4  ;  Pet  =  19-9  ;  CP1  =  157  ;  CPd  =  n-i  ;  Snt  =  3-3. 
The  mouth  is  sub-terminal,  the  lower  jaw  is  curved  without  a  horny  margin  and  both 
anterior  and  posterior  barbels  are  present  although  very  short.  Scales  on  the 
posterior  part  of  the  body  have  parallel  striae  but  are  radiately  striated  on  the 
shoulders  ;  32  scales  in  the  lateral  line,  12  around  the  caudal  peduncle,  5-5  between 
the  dorsal  mid-line  and  the  lateral  line  ;  5-5  between  the  lateral  line  and  the  ventral 
mid-line  and  2-5  between  the  lateral  line  and  the  base  of  the  pelvic  fin. 

The  dorsal  fin  has  four  unbranched  rays,  the  last  is  moderately  thickened  and 
ossified  into  a  smooth  spine,  the  non-articulated  part  of  which  is  n-o  per  cent  of 
the  standard  length.  There  are  nine  branched  rays  and  the  dorsal  margin  of  the 
fin  is  slightly  concave. 

The  anal  fin  has  three  unbranched  and  five  branched  rays.  David  &  Poll 
(1937)  give  six  branched  rays  but  they  were  apparently  misled  by  the  complete 
bifurcation  of  the  last  ray. 

Pharyngeal  bones  and  teeth.  The  right  pharyngeal  bone  is  shown  in  fig.  71  ;  the 
fifth  tooth  in  the  inner  row  is  notable  for  the  presence  of  a  ridge  across  the  concave 
crown.  All  the  teeth  are  hooked. 

There  are  16  short,  broad  gill  rakers  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  Described  by  David  &  Poll  (op.  cit.}  as  dark  grey  on  the  back,  lighter 
grey  below.  Scales  with  a  black  border.  The  colour  now  is  dark  brown  on  the 
back  and  lighter  brown  below. 


88 


K.   E.    BANISTER 


I 


I 


I 


A   REVISION   OF   THE   LARGE   AFRICAN   BARBUS 


FIG.  71.     The  right  pharyngeal  bone  from  the  holotype  of  Barbus  microbarbis. 

DISTRIBUTION.     This  species  is  known  only  from  Lake  Luhondo,  Rwanda. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  microbarbis  is  characterized  by  having  very 
short  barbels  and  a  wide  sub-terminal  mouth  with  a  gently  curving  lower  jaw.  In 
these  two  characters  this  species  approaches  members  of  the  ill-defined  genus 
Varicorhinus  Riippell  but  is  clearly  separate  from  Varicorhinus  ruandae  (see  above) 
with  which  it  is  sympatric. 

It  is  unwise  to  speculate  too  much  on  the  basis  of  one  specimen.  Barbus  micro- 
barbis could  be  an  abnormal  specimen  of  Barbus  altianalis  or  Barbus  acuticeps. 
Groenewald  (1958)  has  shown  how  the  mouth  shape  can  change  markedly.  This 
individual  could  be  a  hybrid  between  a  Barbus  species  and  a  Varicorhinus  species. 

Without  more  specimens  and  a  greater  knowledge  of  the  fauna  of  the  area  the 
relationships  and  nature  of  this  species  must  be  left  in  abeyance. 


Barbus  microterolepis  Boulenger  1902 

Barbus  microterolepis  Blgr.,   1902,  Ann.  Mag.  nat.  Hist.  (7)   10  :  426  ;    Blgr.,   1911,  Cat.  Afr. 
Fish  2  :  23,  fig.  5. 

HOLOTYPE.  A  fish  of  118  mm  S.L.  from  the  Maki  river,  Lake  Zwai,  Ethiopia, 
B.M.  (N.H.)  Reg.  No.  1902.12.13  :  220. 

DESCRIPTION.  The  description  is  based  on  the  only  known  specimen  of  this 
species.  All  measurements  are  expressed  as  percentages  of  the  standard  length. 


K.   E.    BANISTER 


FIG.  72.     Barbus  microterolepis  holotype  (from  Boulenger  191  za). 


D  =  27-1 ;  H  =  25-4 ;  I  =  7-6  ;  IO  =  7-6  ;  MW  =  5-9  ;  Snt  =  7-6  ;  Pet  =  21-2  ; 
CP1  =  16-9  ;  CPd  =  10-2  ;  Ab  =  5-1 ;  Pb  =  6-8. 
The  shape  of  the  body  can  be  seen  in  fig.  72. 

Squamation.  The  lateral  line  has  40  scales.  There  are  7-5  scale  rows  between 
the  dorsal  mid-line  arid  the  lateral  line  and  6-5  scale  rows  between  the  lateral  line 
and  the  ventral  mid-line.  There  are  four  scales  between  the  lateral  line  and  the 
base  of  the  pelvic  fin.  Around  the  caudal  peduncle  there  are  15  or  16  scales.  The 
scales  bear  numerous  fine  parallel  striae. 

Dorsal  fin.  The  dorsal  fin  origin  is  slightly  in  front  of  the  pelvic  fin  insertion.  It 
has  four  unbranched  rays,  the  last  of  which  is  ossified  into  a  smooth  spine  of  length 
22 '9  per  cent.  There  are  eight  branched  rays  the  last  of  which  is  bifid. 

The  anal  fin  has  five  branched  rays  and  three  unbranched  rays. 

Pharyngeal  bones  and  teeth.  The  first  tooth  in  the  inner  row  (fig.  73)  is  small  and 
conical,  the  second  tooth  is  longer  and  thicker  with  a  recurved,  unicuspid  crown. 
The  other  three  teeth  in  this  row  become  progressively  thinner  and  more  hooked. 
There  are  no  molariform  teeth.  The  pharyngeal  tooth  formula  is  2. 3.5. -5. 3. 2. 

Gill  rakers.  Ten  moderately  stout,  slightly  curved  gill  rakers  are  present  on 
the  lower  limb  of  the  first  gill  arch.  Three  gill  rakers  are  present  on  the  upper  limb. 

Coloration.  Boulenger  (op.  cit.)  describes  the  colour  of  the  preserved  specimen 
as  olive  above,  silvery  beneath. 

DISTRIBUTION.  This  species  is  known  only  from  the  Maki  river,  which  is  a  fast- 
flowing  stream  rising  in  the  hills  of  Gouaza  and  emptying  into  Lake  Zwai. 

AFFINITIES.  Any  conclusions  about  this  species,  based  on  one  specimen,  must 
be  regarded  as  tentative.  The  Zwai  basin  contains  two  other  endemic  cyprinids, 
Barbus  ethiopicus  and  Garra  makiensis  (Blgr.  191  la;  Menon  1964).  The  geology 
of  the  area  is  not  known  in  enough  detail  to  be  able  to  decide  whether  the  endemism 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


FIG.  73.     The  left  pharyngeal  bone  from  the  holotype  of  Barbus  microterolepis . 


is  the  result  of  a  long  isolation  or  not.  There  are  several  possibilities  concerning 
Barbus  microterolepis.  It  could  be  an  abnormal  specimen  of  Barbus  intermedius  in 
which  the  number  of  scales  has  been  increased  by  a  chance  mutation.  Barbus 
intermedius  lives  in  the  Zwai  basin  and  differs  from  Barbus  microterolepis  in  the  size 
of  the  scales.  Barbus  microterolepis  could  be  a  valid  species  descended  from  a 
common  ancestry  with  Barbus  intermedius  having  an  increase  in  scale  numbers.  If 
this  is  so  one  must  conclude  that  the  species  must  be  rare  or  inhabiting  inaccessible 
regions  as  only  one  fish  has  ever  been  collected.  A  third  possibility  is  that  Barbus 
microterolepis  is  a  hybrid  between  the  small-scaled  Barbus  ethiopicus  of  Lake  Zwai 
and  Barbus  intermedius.  There  is  no  evidence  at  the  moment  that  Barbus  micro- 
terolepis is  not  a  valid  species  but  more  specimens  are  needed  before  any  firm  con- 
clusions regarding  its  affinities  can  be  reached. 


Barbus  mirabilis  Pappenheim  and  Boulenger  1914 
Barbus  mirabilis  Pappenheim  &  Blgr.,  1914,  Wiss.  Ergebn.  dt.  ZentAfr.  Exped.  Zool.  (3)  5  :  239. 

HOLOTYPE.     A  fish  of  353  mm  S.L.,  Z.M.B.  Reg.  No.  19059. 

DESCRIPTION.  The  description  is  based  on  the  holotype  as  I  have  not  been  able 
to  find  any  other  specimens  which  have  been  referred  to  this  species.  D  =  30-8  ; 
H  =  24-6  ;  1  =  4-2;  IO  =  io-o  ;  MW  =  7-1  ;  Pet  =  19-8  ;  CP1  =  13-8  ; 
CPd  =  1 1 -9  ;  Snt  =  8-5  ;  Ab  =  4-7  ;  Pb  =  5-4.  All  these  measurements  are 
expressed  as  percentages  of  the  standard  length. 


K.   E.    BANISTER 


FIG.  74.     Barbus  mirabilis  holotype. 


The  shape  of  the  body  can  be  seen  in  fig.  74  and  it  presents  a  rather  bulky 
appearance. 

Squamation.  The  striae  on  the  scales  are  slightly  diverging,  less  so  on  the  shoulder 
scales  than  on  the  flanks  or  belly.  There  are  28  scales  in  the  lateral  line  (Pappenheim 
and  Boulenger's  count  of  31  is  the  total  number).  Five  and  a  half  scale  rows  are 
between  the  dorsal  mid-line  and  the  lateral  line  and  the  same  number  between  the 
lateral  line  and  the  ventral  mid-line.  Three  scale  rows  lie  between  the  lateral  line 
and  the  base  of  the  pelvic  fin.  The  scales  on  the  chest  are  reduced  in  size  and  are 
less  well  formed.  There  are  12  scales  around  the  caudal  peduncle. 

The  mouth  is  sub-terminal  and  crescentic  with  a  continuous  lower  lip. 

Dorsal  fin.  Has  four  unbranched  rays.  The  last  ray  is  ossified  into  a  smooth 
spine,  the  non-articulated  part  of  which  is  9-6  per  cent  of  the  standard  length. 
There  are  ten  branched  rays.  The  origin  of  the  dorsal  fin  is  in  front  of  the  pelvic 
fins.  There  is  no  sheath  of  scales  at  the  base. 

Coloration.  The  preserved  fish  is  pale  brown  in  colour  with  a  darker  lower  lobe 
of  the  caudal  and  a  dark  edge  to  the  pectoral  fins.  The  back  and  sides  above  the 
lateral  line  are  slightly  darker  than  the  belly. 

Gill  rakers.     There  are  10  gill  rakers  on  the  lower  arm  of  the  first  gill  arch. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  teeth  (figs.  75  and  76)  have  slightly 
hooked  crowns.  There  is  little  enlargement  of  the  second  tooth  in  the  inner  row. 
The  pharyngeal  tooth  formula  is  2.3. 5. -5.3.2. 

DISTRIBUTION.     Known  only  from  Mawambi  on  the  Ituri  river  (Congo  system). 

AFFINITIES.  In  its  general  shape,  coloration,  striations  of  the  scales,  scale  and 
gill  raker  counts  and  pharyngeal  teeth,  Barbus  mirabilis  closely  resembles  Barbus 
somereni.  The  barbels  are  shorter  than  in  Barbus  somereni  but  the  unique  specimen 
of  Barbus  mirabilis  is  much  bigger  than  the  largest  Barbus  somereni  examined. 
Barbus  somereni  also  usually  lacks  the  dark  edge  to  the  pectoral  fin. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


93 


Without  more  specimens  I  cannot  synonymize  these  two  species  nor  state  their 
relationship  with  any  degree  of  certainty.  It  seems  possible  that,  although  Barbus 
mirabilis  comes  from  the  Congo  system  on  the  other  side  of  the  rift  valley,  it  was 
found  only  some  150  km  from  the  Ruwenzori  mountains  where  Barbus  somereni  is 
common  and  could  represent  a  population  of  Barbus  somereni  that  lived  in  the 
westward-flowing  rivers  of  that  area  and  became  isolated  when  the  rift  valley  formed. 
There  is  a  certain  amount  of  circumstantial  evidence  to  support  this  idea.  Barbus 


FIG.  75.     The  dorsal  aspect  of  the  left  pharyngeal  bone  from  the  holotype  of 

Barbus  mirabilis. 


5mm 


FIG.  76. 


The  lateral  aspect  of  the  left  pharyngeal  bone  from  the  holotype  of 
Barbus  mirabilis. 


94 


K.   E.    BANISTER 


somereni  has  a  wide,  scattered  distribution,  usually  at  high  altitudes  suggesting 
that  it  was  once  a  widespread  species  and  changes  in  the  environment  (whether 
tectonic  or  climatic)  or  competition  have  driven  it  into  higher  altitude  streams. 
What  little  geological  evidence  is  available  (see  under  Barbus  altianalis]  suggests 
that  the  Congo  watershed  extended  further  eastwards  than  it  does  now  and  that  its 
eastern  extremities  were  cut  off  and  their  direction  changed  by  the  rift  valley 
formation.  It  would  therefore  seem  possible  that  an  old-established  large  Barbus 
species  (which  this  must  be  if  the  arguments  have  any  veracity  at  all)  could  be 
found  on  both  sides  of  the  rift  valley.  More  specimens  from  Mawambi  are  badly 
needed  before  this  matter  can  be  pursued  further.  I  have  not  been  able  to  examine 
Pappenheim  &  Boulenger's  (op.  cit.)  unique  specimen  of  Barbus  mawambi  also  from 
Mawambi  but  from  their  description  and  figure  I  am  tempted  to  think  that  this 
species  could  be  the  same  as  Barbus  mirabilis. 


Barbus  oxyrhynchus  Pfeffer  1889 

Barbus  oxyrhynchus  Pfeffer,  1889,  Jb.  Hamb.  Wiss.  Anst.  6  (2)  pi.  8  ;    Pfeffer,  1896,  Thierw. 

O-Afr.  Fische  :  64. 
Barbus  tanensis  Gunther,  1894,  Proc.  zool.  Soc.  Lond.  :    go,  pi.  n  ;    Blgr.,  1911,  Cat.  Afr.  Fish 

2  :  58  ;     Fowler,    1936   (Part),  Proc.   Acad.   nat.   Sci.  Philad.,   88  :  287,   fig.   50   (as  Barbus 

(Lanceabarbus)  tanensis) . 
Barbus  hindii  Blgr.,  1902,  Proc.  zool.  Soc.  Lond.  (2)  :  222,  pi.  16,  fig.  i  ;    Pappenheim  &  Blgr., 

1914,   Wiss.  Ergebn.  dt.  ZentAfr.  Exped.  Zool.   (3)  :  239  ;    Pellegrin,   1909,  Mem.  Soc.  zool. 

Fr.  22  :  281-298. 
Barbus  (Capoeta)  perplexicans  Blgr.,  1902,  Proc.  zool.  Soc.  Lond.  (2)  :  223,  pi.  16,  fig.  2  ;   Pellegrin, 

1909,  Mem.  Soc.  zool.  Fr.  22  :  281-298. 

Barbus  labiatus  Blgr.,  1902,  Proc.  zool.  Soc.  Lond.  (2)  :  223,  pi.  17,  fig.  i. 
Barbus  krapfi  Blgr.,  1911,  Cat.  Afr.  Fish  2  :  54. 
Barbus  mathoiae  Blgr.,  1911,  Cat.  Afr.  Fish  2  :  66. 

Barbus  ahlselli  Lonnberg,  1911,  K.  svenska  Vetensk.  Akad.  Handl.  47  (6)  :  39. 
Barbus  athi  Hubbs,  1918,  Fieldiana  12  (2)  :  9-16. 
^Barbus  babaulti  Pellegrin,  1926,  Bull.  Soc.  zool.  Fr.  51  :  384. 
Barbus  nairobi  Holly,  1928,  Zool.  Anz.  75  (3-4)  :  1-4. 
Barbus  donyensis  Holly,  1929,  Sber.  Akad.  Wiss.  Wien  (4)  :  32-35. 
Varicorhinus  babaulti  :    Berlin  &  Esteve,  1947,  Catalogue  des  Types  des  Poissons  Paris  :  45. 

NOTES  ON  THE  SYNONYMY.  Pfeffer  (1889)  described  Barbus  oxyrhynchus  from 
two  small  specimens  from  the  Rufu  river  (Pangani  system).  Barbus  tanensis  was 
described  from  much  larger  fish  (from  188  to  285  mm  S.L.)  from  the  Thika-Thika 
river  (Tana  system).  Barbus  hindii,  Barbus  perplexicans,  Barbus  labiatus,  Barbus 
krapfi  and  Barbus  mathoiae,  all  came  from  the  Athi,  Tana  and  Pangani  rivers  or  the 
Lorian  swamp,  and  they  were  separated  principally  on  slight  differences  in  propor- 

f  Pellegrin  (1935,  Revue  Zool.  Bot.  afr.  27:  382)  subsequently  described  a  small  Barbus  from  Lake  Kivu 
under  the  name  Barbus  (Agrammobarbus)  babaulti.  Greenwood  (1962)  synonymized  this  species  with 
Barbus  apleurogramma,  Blgr.  191  la.  The  International  rules  of  Zoological  Nomenclature  (Arts.  53  and 
60)  require  one  to  give  a  replacement  name  for  Barbus  (Agrammobarbus)  babaulti,  Pellegrin  1935,  which 
is  here  rejected  as  a  primary  junior  homonym  of  Barbus  babaulti  Pellegrin  1926;  I  propose  that  Pellegrin's 
Lake  Kivu  species  be  called  Barbus  lapsus. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  95 

tion,  especially  of  the  lips,  dorsal  spine  and  body  depth.  The  same  characteristics 
were  used  to  differentiate  the  other  species  from  the  same  localities,  viz.  Barbus 
ahlselli,  Barbus  athi,  Barbus  babaulti,  Barbus  nairobi  and  Barbus  donyensis. 

Fowler  (1936)  was  of  the  opinion  that  the  continued  separation  of  Barbus  hindii, 
Barbus  tanensis  and  Barbus  mathoiae  was  unjustified.  Allowing  for  the  known 
vagaries  of  lip  development  (Groenewald  1958)  and  for  the  allometry  of  various 
parts  of  the  body  with  growth  it  seems  reasonable  to  conclude  that  all  these  nominal 
species  from  a  few  adjacent  rivers  are  preferable  to  the  same  species.  Certainly 
the  adults  have  the  same  general  appearance  and,  as  the  histograms  indicate,  normal 
variation  (i.e.  Poisson  distribution)  of  various  characters  is  present  when  large  series 
of  specimens  are  examined.  The  only  species  over  which  I  have  any  doubts  is 
Barbus  labiatus  (known  only  from  two  specimens)  which  has  a  shallower  body  and  a 
slightly  longer  head  than  the  others.  But  with  only  two  specimens  available  I 
cannot  attribute  too  much  significance  to  slight  differences. 

It  has  not  proved  possible,  using  the  characters  considered  useful  in  this  paper, 
to  divide  Barbus  oxyrhynchus  into  smaller  groups.  This  is  only  to  be  expected  as 
the  overall  range  of  morphometric  characters  is  not  very  great  and  the  head  waters 
of  the  Athi  and  Tana  systems  are  extremely  close. 

One  fish  (Coll.  M.A.C.T.  No.  47341)  from  the  Malagarasi  river  was  attributed  to 
Barbus  krapfi  by  David  (1937)  and  to  Barbus  lestradei  (=  Barbus  caudovittatus]  by 
Poll  (1953).  Barbus  oxyrhynchus  has  not  been  found  in  the  Malagarasi,  yet  I 
cannot  easily  place  this  specimen  with  any  of  the  Malagarasi  species.  The  pectoral 
fin  is  slightly  shorter  than  in  Barbus  oxyrhynchus,  but  in  other  features  it  corresponds 
well.  I  can  only  assume  either  that  Barbus  oxyrhynchus  is  found  in  the  Malagarasi 
but  is  extremely  rare  or  that  this  specimen  is  a  deformed  member  of  another  species 
if  it  is  not  a  new  species. 

A  specimen  in  the  Stockholm  Museum,  No.  8061,  identified  as  Barbus  percivali 
Blgr.  ( =  Barbus  neumayeri  Fischer,  fide  Greenwood  1962  :  178)  from  Kibonoto  on 
the  Sanya  river,  belongs  to  Barbus  oxyrhynchus. 

There  are  two  fish  from  the  Ruaha  river  (Rufiji  system)  in  the  collection  of  the 
Central  Africa  Museum,  Tervuren,  which  belong  possibly  to  this  species.  The 
Rufiji  is  to  the  south  of  the  Pangani  and  its  fauna  is  very  poorly  known.  These  two 
fish  are  extremely  deep  bodied,  their  standard  lengths  are  156  and  164  mm  with 
body  depths  of  38-4  and  40  per  cent  respectively.  Their  other  features  are  within 
the  Barbus  oxyrhynchus  range  except  for  the  dorsal  spines  which  are  short  and 
articulated  distally  (respectively  11-5  and  15-2  per  cent).  Although  at  the  moment, 
these  specimens  are  considered  as  belonging  to  this  species  they  are  not  included  in 
the  data  given  in  the  description  below.  Further  specimens  may  show  that  the 
Rufiji  fishes  are  of  a  different  species  or  are  a  discrete  population  of  Barbus  oxyrhyn- 
chus. Pappenheim  &  Boulenger  (1914)  record  Barbus  hindii  from  the  Ituri  river 
but  the  very  low  lateral  line  count  would  seem  to  refer  these  fish  to  Barbus 
mawambiensis. 

LECTOTYPE.  The  lectotype,  a  fish  of  48  mm  S.L.  from  the  Pangani  river,  Ham- 
burg Museum,  No.  11339,  was  selected  by  Ladiges  et  alii  (1958).  I  have  not  been 
able  to  examine  this  specimen  but  it  was  compared  for  me  by  Dr  W.  Ladiges  who 


96 


K.    E.    BANISTER 


FIG.  77.     Barbus  oxyrhynchus  paralectotype. 


kindly  sent  me  the  largest  paralectotype  (40  mm  S.L.)  which  is  figured  above 
(fig-  77)- 

DESCRIPTION.  The  description  is  based  on  108  specimens,  including,  in  addition 
to  the  types  of  Barbus  oxyrhynchus,  the  types  of  Barbus  tanensis,  Barbus  hindii, 
Barbus  perplexicans,  Barbus  labiatus,  Barbus  krapfi,  Barbus  mathoiae,  Barbus 
ahlselli,  Barbus  babaulti  and  Barbus  athi.  The  size  range  of  the  specimens  is  from 
28  to  369  mm  S.L.  Because  the  lectotype  is  a  juvenile  fish  figs.  78,  79  and  80  show 
larger  specimens  indicating  the  variations  produced  by  growth  in  this  species. 

The  morphometric  data  are  expressed  in  tabular  form  as  follows  and  unless 
otherwise  stated  are  expressed  as  a  percentage  of  the  standard  length. 


D 

H 

I 

IO 

MW 

Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


26-9 

7-6 

8-9 

6-4 

24-4 

16-9 

12-8 

8-5 

5-9 

7-1 


s.d. 
2-8 

2-5 
1-9 
i-i 
0-9 
1-9 
1-4 
i-i 
0-9 
1-4 
1-4 


0-2 
O-2 
O-I 
O-I 
O-2 
O'l 
O'l 
O'l 
O-I 
O'l 


range 

26-1-39-4 

23-0-32-6 

4-4-n-i 

6-9-12-5 

4-3-  8-7 

21-3-28-6 

14-1-20-0 

10-8-15-5 

6-3-11-0 

3-1-10-7 

3-6-12-4 


The  standard  deviation  is  higher  than  for  many  species  described  in  this  paper 
because  of  the  size  range  and  marked  allometry  of  the  specimens  examined.  This 
is  especially  noticeable  in  the  body  depth  ;  in  fishes  of  less  than  100  mm  S.L. 
(measurements  expressed  as  a  percentage  of  the  standard  length  unless  otherwise 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


97 


FIG.  78.     Barbus  oxyrhynchus  (type  of  B.  labiatus  from  Boulenger  191  la). 


FIG.  79.     Barbus  oxyrhynchus  (type  of  B.  hindii  from  Boulenger  191  xa). 


stated),  x  =  29-3  ;  s.d.  =  2-2  ;  s.e.  =  0-4 ;  range  =  26-1-35-0 ;  the  upper  end  of 
this  range  is  extended  by  two  small  fishes  from  the  Athi  river  which  have  a  pro- 
nounced keel  leading  up  to  the  dorsal  fin.  Fishes  of  S.L.  101-170  mm,  x  =  32-2  ; 
s.d.  =  1-6  ;  s.e.  =  0-2  ;  range  =  29-5-35-8.  Fishes  of  S.L.  above  171  mm, 
x  =  33'6  ;  s.d.  =  2-7  ;  s.e.  =  0-5  ;  range  =  28-4-39-4.  In  this  case  the  lower  end 
of  the  range  is  extended  by  the  shallow-bodied  specimens  previously  referred  to 
Barbus  labiatus  (see  below). 

The  typical  adult  fish  has  a  fairly  flat  ventral  profile.  The  dorsal  profile  rises 
sharply  from  the  snout  to  the  origin  of  the  dorsal  fin  after  which  it  follows  a  sloping 
concave  course  down  to  the  caudal  fin.  The  same  type  of  body  shape  is  shown  by 
the  two  specimens  attributed  to  Barbus  labiatus  except  that  the  body  is  less  deep. 
The  snout  is  obtusely  pointed.  The  mouth  ranges  from  ventral  and  curved  in 
most  specimens  through  the  wide,  cutting  mouth  of  the  types  of  Barbus  perplexicans 
to  the  rubber  lips  of  the  type  of  Barbus  labiatus.  The  mean  length  for  the  pectoral 
fin  is  sufficiently  great  for  it  to  serve  as  a  diagnostic  character  for  the  species. 


98 


K.   E.   BANISTER 


FIG.  80.     Barbus  oxyrhynchus  (type  of  B.  krapfi  from  Boulenger  191  la). 


FIG.  81.     A  scale  from  a  specimen  of  Barbus  oxyrhynchus  (the  type  of  B.  mathoiae) 
to  show  the  distribution  of  the  striae. 


The  caudal  peduncle  is  compressed,  short  and  deep  with  a  mean  length/depth 
ratio  of  1-32. 

Squamation.  The  scales  bear  many  parallel  or  slightly  converging  striae  (fig.  81). 
The  lateral  line  count  is  low,  from  21  to  28  scales;  21  (f.i),  22  (f.io),  23  (f.J-5), 
24  (f-36),  25  (f.3o),  26  (f.8),  27  (f.6),  28  (f.i).  The  type  of  Barbus  labiatus  has 
24  on  one  side  and  27  on  the  other.  The  lateral  line  follows  a  rather  dipping  course. 
There  are  4-5  (f.ioo)  rarely  3-5  (f.6)  or  5-5  (f.2)  scales  between  the  dorsal  mid-line 
and  the  lateral  line  and  4-5  (f.8i)  rarely  3-5  (f.i6)  or  5-5  (f.3)  scales  between  the 
lateral  line  and  the  ventral  mid-line.  This  count  was  unobtainable  from  some 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  99 

specimens.  There  are  2-5  (f.65),  2  (f./j.)  or  1-5  (f-32)  scales  between  the  lateral  line 
and  the  base  of  the  pelvic  fin.  Again  this  count  was  not  possible  in  some  specimens. 
Around  the  caudal  peduncle  there  are  12  (1.99)  or  n  (f.5)  scales  ;  this  count  too 
was  unobtainable  for  some  specimens. 

Dorsal  fin.  There  are  four  unbranched  rays.  The  last  of  these  is  ossified  into  a 
long,  moderately  broad,  straight  or  gently  curving  spine.  This  fourth  ray  has  a 
mean  length  of  25-3  with  a  range  from  16-6  to  35-7  per  cent,  s.d.  =  4-6,  s.e.  =  0-5. 

There  are  from  eight  to  ten  branched  rays :  8  (1.5),  9  (f.82),  10  (f.2i) ;  the  types 
of  Barbus  oxyrhynchus  are  somewhat  atypical  in  having  only  eight  branched  rays. 
The  dorsal  margin  is  strongly  concave  and  the  base  of  the  dorsal  fin  frequently  has 
an  enveloping  sheath  of  scales. 

The  anal  fin  has  three  unbranched  rays  and  five  (very  rarely  six)  branched  rays. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  tooth  formula  is  constant  at 
2. 3. 5. -5. 3. 2.  The  first  tooth  in  the  inner  row  is  tall,  thin  and  has  a  slightly  hooked 
crown.  It  is  slightly  angled  towards  the  second  tooth.  This  tooth  is  longer  than 
wide  (fig.  83)  and  again  hooked  distally.  Teeth  3,  4  and  5  become  progressively 
thinner  and  develop  articular  surfaces  on  the  posterior  face  of  the  crown.  The 
fifth  tooth  is  distinctly  recurved  but  the  terminal  hook  remains.  The  teeth  of  the 
second  and  third  rows  resemble  the  fifth  tooth  of  the  ventral  row  in  general  shape 
but  are  much  smaller. 


FIG.  82.     Scatter  diagram  of  the  dorsal  spine  length  as  a  percentage  of  the  standard  length 
against  the  standard  length  for  the  entire  sample  of  Barbus  oxyrhynchus. 


K.   E.    BANISTER 


FIG.  83.     The  left  pharyngeal  bone  from  Barbus  oxyrhynchus  (a  syntype  of  B.  tanensis). 


The  gill  rakers  range  from  n  to  16  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  Copley  (1958)  states  that  the  colour  of  Barbus  tanensis  varies  in  live 
fish  from  olive-green  to  deep  olive-brown  on  the  back  shading  to  silver  on  the  belly. 
Fowler  (1936)  figures  Barbus  tanensis  with  a  dark  spot  on  the  caudal  peduncle  ;  I 
have  not  seen  this  in  any  of  the  specimens  I  examined.  The  colour  in  alcohol  is 
sandy-brown  on  the  back  shading  to  a  pale  brown  to  silver  on  the  belly.  The  scale 
margins  are  frequently  a  deeper  shade  of  brown  than  the  centre  of  the  scale. 

DISTRIBUTION.  Specimens  are  known  from  the  Rufu  river,  Pangani  system  ; 
the  Kibwesi,  Makindu,  Tsavo,  Regati,  Mathoiya  and  Thika-Thika  rivers,  Athi-Tana 
systems  ;  the  Rufiji  river ;  Nero-Narok  system,  Lorian  swamp  (in  the  Northern 
Euasso  Nyiro  system).  This  species  is  widespread  throughout  the  Athi  and  Tana 
systems  (Copley  1958). 

The  locality  G.  el  Narua  given  in  Boulenger  (191  la :  57)  is  the  subject  of 
some  dispute  and  is  discussed  in  detail  on  p.  71. 

I  have  not  had  the  opportunity  to  examine  any  Barbus  specimens  from  Mozambique 
north  of  the  Zambezi  so  it  is  possible  that  the  range  of  Barbus  oxyrhynchus  may 
extend  further  south. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  oxyrhynchus  is  closely  related  to  Barbus 
bynni,  Barbus  gananensis  and  Barbus  longifilis  and  the  points  of  differences  between 
Barbus  oxyrhynchus  and  the  three  other  species  will  be  found  on  pp.  36  and  129. 

Comments  on  clinal  phenomena  within  this  group  of  species  and  exemplified  by 
Barbus  oxyrhynchus  are  found  on  p.  129. 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  101 

Barbus  pagenstecheri  Fischer  1884 

Barbus  pagenstecheri  Fischer,  1884,  Jb.  hamb.  wiss.  Anst.  1  :  30  ;    Pfeffer,  1896,  Thierw.  O.-Afr. 
Fische  5  :  65  ;    Boulenger,  1911,  Cat.  Afr.  Fish  2  :  72. 

LECTOTYPE.  The  original  description  was  based  on  two  specimens  (Nos.  H34I 
and  H342  in  the  Hamburg  Museum  of  315  and  217  mm  S.L.  respectively).  Boulen- 
ger (191  la)  redescribed  the  species  on  the  basis  of  the  smaller  specimen  and  implied 
that  this  specimen  was  the  lectotype  (he  used  the  word  '  type '  and  mentioned  that 
a  larger  specimen  had  also  been  referred  to  this  species).  Ladiges  et  alii  (1958), 
in  a  non-revisional  work,  designated  the  larger  specimen  (H34i)  as  the  lectotype. 
Here  I  follow  Boulenger  in  accepting  the  smaller  specimen  as  the  lectotype  and 
regarding  the  larger  specimen  as  the  paralectotype. 

DESCRIPTION.  The  description  is  based  on  the  only  two  specimens  referred  to 
this  species.  The  morphometric  data  are  shown  below  and  in  each  case  the  lectotype 
comes  first.  L  =  217,  315  mm  ;  D  =  26-3,  29-8  ;  H  =  24-0,  30-8  ;  I  =  6-4,  4-8  ; 
IO  =  7-4,  10-2  ;  MW  =  6-9,  8-9  ;  Pet  =  20-3,  24-8  ;  CP1  =  17-5,  17-5  ;  CPd 
=  io-i,  12-3  ;  Snt  =  6-9,  9-8  ;  Ab  =  4-1,  5-1  ;  Pb  =  5-2,  6-4. 

Unless  otherwise  stated  all  measurements  are  expressed  as  percentages  of  the 
standard  length. 


5cm 


FIG.  84.     Barbus  pagenenstecheri  lectotype. 

The  body  (fig.  84)  is  elongated,  the  dorsal  profile  rises  gradually  and  smoothly 
from  the  snout  to  the  origin  of  the  dorsal  fin.  The  caudal  peduncle  is  about  half  as 
long  again  as  it  is  deep.  The  mouth  is  sub-terminal  with  a  sharp  edge  to  the  lower 
jaw  in  the  lectotype  but  rubber  lips  are  developed  in  the  paralectotype.  The  snout 
is  blunter  in  the  lectotype  than  in  the  paralectotype.  Both  specimens  have  a  clearly 
defined  rostral  groove.  Numerous  small  tubercules  are  present  on  the  snout  and 
the  cheeks  of  both  specimens. 

Squamation.  There  are  26  or  27  scales  in  the  lateral  line  series,  4-5  (f.2)  scales 
between  the  dorsal  mid-line  and  the  lateral  line,  4-5  (f.2)  scales  between  the  lateral 
line  and  the  ventral  mid-line  and  2-5  (f.2)  scales  between  the  lateral  line  and  the 


IO2 


K.   E.    BANISTER 


5  mm 


FIG.  85.     Barbus  pagenenstecheri  -  dorsal  and  lateral  views  of  the  left  pharyngeal  bone. 


base  of  the  pelvic  fin.     Twelve  scales  encircle  the  caudal  peduncle.     The  exposed 
parts  of  the  scales  bear  numerous,  sinuous,  more  or  less  parallel  striae. 

Dorsal  fin.  There  are  four  unbranched  rays  and  eight  branched  rays  in  the  dorsal 
fin.  The  last  unbranched  ray  is  weakly  ossified  into  a  smooth  spine,  14-3,  13-7  per 
cent  S.L.  The  dorsal  margin  of  the  fin  is  slightly  concave.  The  dorsal  fin  origin 
is  slightly  in  advance  of  the  pelvic  fin  origin. 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays. 

Pharyngeal  bones  and  teeth.  Pharyngeal  bones  are  present  only  in  the  paralecto- 
type.  The  pharyngeal  teeth  number  2.3.5. -5.3. 2.  The  teeth  are  widely  spaced 
(fig.  85).  The  lateral  flange  is  confined  to  the  angle  ot  the  bone  level  with  the 
fourth  and  fifth  teeth  of  the  inner  row. 

Gill  rakers.     There  are  12  or  15  gill  rakers  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  The  colour  of  alcohol-preserved  specimens  is  dark  brown  on  the  back 
paling  to  yellow-brown  on  the  ventral  surface.  The  edges  of  the  scales  on  the  back 
and  dorsal  part  of  the  flanks  are  lighter  than  the  centres.  The  fins  are  dark  grey- 
brown. 

DISTRIBUTION.  Fischer  (1884)  collected  the  fish  from  a  stream  flowing  down 
Mount  Kilimanjaro.  Bailey  (1969)  gives  the  locality  as  in  the  Pangani  system. 
There  is  no  evidence  for  this  because  only  the  streams  on  the  south  face  of  Mount 
Kilimanjaro  flow  into  the  Pangani.  The  streams  on  the  eastern  face  flow  into  the 
Tsavo  river  whilst  the  streams  on  the  northern  and  western  faces  have  no  outlet 
to  the  sea.  All  of  the  streams  are  covered  by  the  locality  description  '  German  East 
Africa'.  Detailed  information  on  the  expeditions'  collecting  sites  is  lacking. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  pagenstecheri  is  a  species  of  uncertain  affini- 
ties, its  distribution  is  localized  and  it  is  surrounded  by  waters  containing  Barbus 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  103 

oxyrhynchus.  I  have  not  seen  any  specimens,  nor  have  I  been  able  to  find  any 
reliable  records,  of  Barbus  oxyrhynchus  from  the  Kilimanjaro  region  and  it  is  there- 
fore possible  that  the  two  specimens  referred  to  Barbus  pagenstecheri  may  represent 
a  local  population  of  Barbus  oxyrhynchus.  The  differences  between  these  two  fishes 
and  equal-sized  specimens  of  Barbus  oxyrhynchus  are  marked.  The  former  fishes 
have  shallow  bodies  and  weak  dorsal  spines  whilst  the  latter  have  deeper  bodies 
and  strong  dorsal  spines.  The  lateral  line  scale  count  in  Barbus  pagenstecheri  is 
at  the  upper  end  of  the  Barbus  oxyrhynchus  range  and  the  scale  striations  are  similar 
in  both  species.  Barbus  mariae  from  the  Athi  river  can  easily  be  distinguished  from 
Barbus  pagenstecheri  by  its  'rhinoceros  horn'  (see  p.  84).  With  only  two  specimens 
available  it  is,  I  feel,  preferable  to  maintain  the  species  Barbus  pagenstecheri  rather 
than  to  regard  these  specimens  as  local  variants  of  Barbus  oxyrhynchus.  Further 
collections  from  the  Kilimanjaro  region  are  needed  to  reach  a  satisfactory  con- 
clusion. 


Barbus  paucisquamatus  Pellegrin  1935 

Barbus  altianalis  var.  labiosa  (part)   Pellegrin,   1933,   Bull.  Soc.  zool.  Fr.  58  :  169   (only  the 

paratype,  from  Loama). 

Barbus  altianalis  var.  paucisquamata  Pellegrin,  1935,  Revue  Zool.  Bot.  afr.  27  (3)  :  378. 
Barbus  altianalis  var.  longifilis  (part)  Pellegrin,  1935,  Revue  Zool.  Bot.  afr.  27  (3)  :  378  (only 

the  specimen  from  Nyabarongo). 
Barbus  altianalis  var.  lobogenysoides  Pellegrin,  1935,  Revue  Zool.  Bot.  afr.  27  (3)  :  380. 

NOTES  ON  THE  SYNONYMY.  The  specimens  of  the  varieties  of  Barbus  altianalis 
described  by  Pellegrin  (op.  cit.)  which  came  from  rivers  in  the  Congo  system  consti- 
tute a  species  distinct  from  Barbus  altianalis. 

The  name  labiosa,  although  the  oldest  infra-specific  name,  is  not  available  for  this 
newly  defined  species  as  the  holotype  of  that  variety  (a  fish  from  Lake  Kivu)  is  a 
specimen  of  Barbus  altianalis  altianalis.  The  type  series  of  Barbus  altianalis  var. 
paucisquamata  is  homogeneous  and  the  infra-specific  name  paucisquamata  (when 
the  gender  is  changed  to  form  paucisquamatus  -  Article  30)  is  available  and  is  used 
accordingly. 

There  are  certain  inaccuracies  in  the  original  description  of  the  specimens  of 
Barbus  altianalis  var.  paucisquamata.  Pellegrin  (1935  :  379)  lists  three  specimens 
of  lengths  130  +  35  =  165,  140  +  40  =  180,  130  +  35  =  165.  The  measurements 
presumably  refer  to  the  standard  length,  '  tail  fin '  length  and  the  total  length.  The 
three  syntypes  (Paris  Museum  Nos.  35-76,  35-77,  35-78  are  of  124,  164  and  116  mm 
S.L.  respectively.  M.A.C.T.  specimen  42932  is  registered  as  a  'co-type',  there 
is  a  label  with  this  fish  saying  'co-type,  don.  de  Mus.  Paris  Loc.  Kivu,  rec.  Guy 
Babault'.  This  fish  is  of  177  mm  S.L. 

LECTOTYPE.  Specimen  35-76  in  the  Paris  Museum  is  designated  the  lectotype. 
This  specimen  of  124  mm  S.L.  is  closest  in  size  to  any  of  the  Pellegrin  measurements. 
It  is  also  the  only  specimen  with  a  precise  locality  which  is  Kitembo,  Nyabarongo 
river. 


io4 


K.   E.    BANISTER 


d 

fi 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  105 

DESCRIPTION.     The  description  is  based  on  12  specimens  from  80  to  248  mm  S.L. 

x  s.d.  s.e.  range 

L  80    -248  mm 

D  26-8  1-6  0-5  24-0-30-0 

H  25-5  1-3  0-4  23-8-27-6 

I  7-2  1-2  0-3                      5-4-  9-4 

IO  7-6  0-9  0-2                      6-3-  8-5 

MW  5-8  0-4  o-i                      5-2-  6-2 

Pet  21-2  0-9  0-3  2O-2-22-8 

CP1  18-0  1-2  0-3  i6'4-i9'7 

CPd  ii-o  i-o  0-3  9-4-13-0 

Snt  8-0  0-9  0-3  6-8-  9-3 

Ab  7-5  1-4  0-4  5-8-10-6 

Pb  8-6  1-7  0-4  6-2-11-2 

All  measurements  are  expressed  as  percentages  of  the  standard  length.  The  body 
is  compressed.  The  dorsal  profile  of  the  snout  is  curved,  the  ventral  profile  is  flat. 
The  mouth  is  ventral  and  horse-shoe  shaped.  The  lower  lip  is  well  defined  and 
varies  between  continuous  with  a  slight  mental  lobe  and  medially  discontinuous. 
The  upper  lip  forms  the  ventral  edge  of  the  snout.  The  barbels  are  conspicuously 
long. 

Squamation.  The  lateral  line  has  27  (f.3),  28  (f.4)  or  29  (f-5)  scales.  Between 
the  dorsal  mid-line  and  the  lateral  line  there  are  4-5  (1.12)  scales.  Only  seven 
specimens  were  in  sufficiently  good  condition  for  the  scales  between  the  lateral  line 
and  the  ventral  mid-line  to  be  counted  ;  five  specimens  have  4-5  scales  and  two 
specimens  have  5-5  scales.  Two  and  a  half  scales  are  present  between  the  lateral 
line  and  the  pelvic  fin  insertion.  There  are  12  scales  around  the  caudal  peduncle. 

The  striae  on  the  scales  are  numerous  and  more  or  less  parallel.  The  striae  of 
scales  on  the  upper  part  of  the  body  tend  to  diverge  slightly  whilst  scales  on  the 
lower  part  of  the  body  tend  to  have  striae  that  are  parallel  or  slightly  converging. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays,  the  last  one  is  ossified  into  a 
smooth,  straight  or  slightly  curved  spine  (x  =  19-7  ;  s.d.  =  2-7  ;  s.e.  =  0-8  ; 
range  14-1-22-5).  There  are  nine  (f.n)  or  eight  (f.i)  branched  rays.  The  dorsal 
margin  of  the  fin  is  only  very  slightly  concave.  The  origin  of  the  dorsal  fin  is  in 
front  of  the  origin  of  the  pelvic  fins. 

Pharyngeal  bones  and  teeth.  No  exceptions  to  the  pharyngeal  tooth  formula 
2.3.5. -5.3. 2.  were  found.  The  teeth  are  shown  in  fig.  87.  The  anterior  edentulous 
process  has  its  distal  half  at  an  angle  to  the  proximal  half. 

Gill  rakers.  There  are  n  (f.3)  or  10  (f.3)  on  the  lower  limb  of  the  first  gill  arch 
in  the  six  specimens  examined. 

Coloration.  All  the  preserved  specimens  are  dark  fish.  Dark-brown  pigment  is 
present  on  the  distal  parts  of  the  dorsal,  anal,  pelvic  and  pectoral  fins.  On  the 
caudal  fin  the  pigment  is  concentrated  at  the  margins. 


io6 


K.   E.    BANISTER 


FIG.  87.     Barbus  paucisquamatus  -  left  pharyngeal  bone  from  the  lectotype. 

The  body  is  dark  brown  but  the  scales  have  a  noticeably  high  reflectivity.  The 
skin  of  the  cheeks  and  snout  has  a  silvery  layer  below  the  brown  pigment.  The 
dorso-lateral  surface  of  both  barbels,  particularly  the  anterior  barbel,  is  pigmented. 

DISTRIBUTION.  Known  from  the  Loama  river,  the  Nyabarongo  river  and  the 
Luhoho  Congo. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  paucisquamatus  is  a  fish  with  27-29  scales 
in  the  lateral  line  series,  dark  pigment  widespread  over  the  body  but  not  obscuring 
a  very  silvery  layer  beneath  and  scales  on  which  the  striae  vary  with  the  position  of 
the  scale  on  the  body. 

Its  affinities  are  uncertain  as  there  is  an  absence  of  clear-cut  similarities  or  dis- 
tinctions with  other  species.  It  is  sympatric  with  Barbus  longifilis  from  which  it 
can  be  distinguished  by  its  shorter  barbels,  shallower  body  and  weaker  dorsal  spine. 

Although  Barbus  paucisquamatus  was  described  as  a  variety  of  Barbus  altianalis, 
the  caudal  peduncle  is  conspicuously  longer  and  shallower  than  in  that  species, 
and  the  overall  impression  gained  from  the  body  shape  and  colour  is  that  there  is  no 
close  relationship  between  Barbus  altianalis  and  Barbus  paucisquamatus. 

The  heavy  pigmentation  and  a  similar  mouth  are  also  found  in  Barbus  caudo- 
vittatus  (p.  40)  but  at  the  moment  I  do  not  have  enough  information  to  comment 
further  on  the  possibilities  of  a  relationship  between  these  two  species,  nor  between 
Barbus  paucisquamatus  and  the  morphologically  somewhat  similar  Barbus  trachy- 
pterus. 


Barbus  platyrhinus  Boulenger  1900 

Barbus  platyrhinus  Blgr.,  1900,  Ann.  Mag.  nat.  Hist.  (7)  6  :  479. 

HOLOTYPE.     A  fish  of  347  mm  S.L.  from  Usamburu,  Lake  Tanganyika,  B.M. 
(N.H.)  Reg.  No.  1906.9.6  :  12. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


107 


DESCRIPTION.     Based  upon  seven  specimens,  the  holotype  and  six  fishes  M.A.C.T. 
Nos.  89789-92,  130658-60. 


L 

D 

H 

I 

IO 

MW 

Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


29-6 

257 

7-2 

9-8 

6-7 

21-4 

16-5 

12-2 

8-8 
3'4 
4'5 


s.d. 
2-4 

I-O 

0-9 
0-9 
0-9 

i'3 
i-i 

0-8 
0-6 
0-7 


s.e. 

0-9 
0-4 
0-4 


0-4 


0-2 


range 

57    -347mm 

26-1-32-3 

24-2-27-2 

6-3-  9-1 

8-2-10-9 

5-7-  8-1 
19-6-23-7 
15-2-18-7 
10-9-13-7 

7-6-  9-5 

2-1-  4-3 


Barbus  platyrhinus  is  a  heavy-bodied  fish,  the  bulky  appearance  becoming  more 
exaggerated  in  larger  specimens.  The  mouth  is  ventral  and  the  snout  has  a  slightly 
bulbous  profile  above  the  upper  jaw. 


FIG.  88.     Barbus  platyrhinus  holotype  (from  Boulenger  191  la). 


Squamation.  The  lateral  line  follows  a  fairly  straight  course.  The  dip  in  the 
middle  of  the  line  is  shallow.  The  lateral  line  scale  counts  are  38  (f.2),  39  (1.3), 
40  (f.i),  41  (f.i).  There  are  6-5  (f.i)  or  7-5  (f.6)  scales  between  the  dorsal  mid-line 
and  the  lateral  line.  There  are  6-5  (f.i),  7-5  (f.2),  8-5  (f.4)  scales  between  the 
lateral  line  and  the  ventral  mid-line.  There  are  3-5  (f.2)  or  4-5  (f.4)  scales  between 
the  lateral  line  and  the  base  of  the  pelvic  fin.  Sixteen  (f.3)  or  18  (f.4)  scales  encircle 
the  caudal  peduncle,  the  scales  are  longitudinally  striated. 

Dorsal  fin.  The  dorsal  fin  has  four  unbranched  rays.  The  last  unbranched  ray 
ranges  from  10-1  to  24-6  per  cent  S.L.  with  a  mean  value  of  18-1  per  cent.  There 
are  8  (f.i),  9  (f.5)  or  10  (f.i)  branched  rays.  The  origin  of  the  dorsal  fin  is  slightly 
in  advance  of  the  origin  of  the  pelvic  fins. 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays. 


io8 


K.    E.    BANISTER 


FIG.  89.     The  pharyngeal  bones  from  the  holotype  of  Barbus  platyrhinus. 


Pharyngeal  bones  and  teeth.  The  only  pharyngeal  bones  that  I  have  been  able  to 
examine  are  from  the  holotype  and  are  apparently  aberrant  (fig.  89)  ;  the  pharyngeal 
formula  is  4.3.2.  The  inner  row  lacks  the  first  tooth  present  in  other  species,  the 
first  tooth  present  in  Barbus  platyrhinus  is  the  homologue  of  the  usual  second  tooth 
and  is  referred  to  as  the  second  tooth.  The  second  tooth  is  large  and  molariform. 
The  third  tooth  has  a  rounded  crown  wider  than  long  with  a  slight  concavity  in 
the  posterior  face  of  the  left-hand  tooth  and  a  deep  concavity  in  the  anterior  face 
of  the  right-hand  tooth.  Jhe  fourth  tooth  is  smaller  than  the  preceding,  the 
anterior  edge  of  the  crown  is  its  highest  point  and  the  posterior  face  is  concave. 
This  tooth  is  wider  than  it  is  long.  The  fifth  tooth  is  slight  and  its  spatulate  crown 
is  very  close  to  the  preceding  crown.  On  the  left-hand  bone  the  first  tooth  of  the 
middle  row  is  missing  but  the  scar  is  present.  Its  counterpart  is  mamilliform  and 
wider  than  long.  It  is  physically  closer  to  the  anterior  tooth  of  the  dorsal  row  than 
to  the  second  tooth  of  the  middle  row.  The  second  and  third  teeth  become  pro- 
gressively smaller.  They  are  complete  on  the  left-hand  bone  but  either  aberrant 
or  in  the  process  of  being  replaced  on  the  right-hand  bone.  The  two  small  teeth  in 
the  outer  row  are  wider  than  long  with  a  concave  posterior  face. 

The  pharyngeal  bones  themselves  are  short  and  thick. 

There  are  between  n  and  13  broad,  bilobed  gill  rakers  on  the  lower  part  of  the 
first  gill  arch. 

DISTRIBUTION.  This  species  has  been  recorded  at  Usamburu  and  Uvira,  Lake 
Tanganyika  and  from  the  Koki  river,  an  affluent  of  Lake  Tanganyika.  Poll  (1953) 
believes  that  it  is  rare  in  the  lake  and  may  be  commoner  in  the  affluent  rivers. 

DIAGNOSIS  AND  AFFINITIES.  This  species  is  sympatric  with  Barbus  tropidolepis 
to  which,  at  the  very  least,  it  is  closely  related.  Both  species  have  a  high  number 
of  lateral  line  scales  and  similarly  shaped  pharyngeal  teeth.  The  major  difference 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  109 

between  them  is  the  development  of  the  fatty  ridges  on  the  scales  in  Barbus  tropido- 
lepis  and  the  universal  presence  of  barbels  in  Barbus  platyrhinus.  A  larger  series  of 
specimens  of  this  species  and  some  information  on  their  ecology  may  show  that  they 
are  not  a  valid  species,  but  for  the  moment  it  must  be  retained  as  there  is,  in  some 
characters,  no  overlap.  I  have  not  yet  found  any  fish  which  cannot  with  certainty 
be  placed  in  either  Barbus  platyrhinus  or  Barbus  tropidolepis ,  but  the  specimens 
examined  have  been  few  in  number.  The  possibility  that  Barbus  platyrhinus  is  a 
hybrid  must  also  be  borne  in  mind. 


Barbus  ruasae  Pappenheim  and  Boulenger  1914 

Barbus  ruasae  Pappenheim  and  Blgr.,  1914,  Wiss.  Ergebn.  dt.  ZentAfr.  Exped.  Zool.  (3)  5  :  238, 
pi.  2,  fig.  2. 

LECTOTYPE.  The  lectotype  is  the  smaller  of  the  two  type  specimens  of  Pappen- 
heim &  Boulenger,  a  fish  of  155  mm  total  length  and  126  mm  S.L.  This  is  the 
specimen  figured  by  the  authors.  The  lectotype  and  paralectotype  (a  fish  of 
128  mm  S.L.)  are  in  the  Zoological  Museum,  Berlin,  both  registered  as  ZMB  19053. 
The  locality  is  Mkunga  in  Rwanda. 

DESCRIPTION.  The  description  is  based  on  five  specimens,  the  two  from  Mkunga 
(see  above)  and  three  specimens,  M.A.C.T.  Nos.  91755,  91756,  92214,  from  the 
Nyabugogo,  an  affluent  of  the  Lusine  about  64  km  SSE  of  the  type  locality. 

x  s.d.  s.e.  range 

L  103  -135  mm 

D  25-6  1-6  0-7  23-7-28-1 

H  27-0  0-9  0-4  25-8-28-1 

I  8-6  0-6  0-3  8-0-  9-7 

IO  7-3  0-3  o-i  7-0-  7-8 

MW  6-5  i-o  0-4  5-6-  8-0 

Pet  21-8  2-2  I-O  18-3-23-4 

CP1  16-2  1-3  0-6  14-7-18-4 

CPd  10-6  0-5  0-2  10-1-11-5 

Snt  8-4  0-5  0-2  7-8-  8-9 

Ab  5-3  1-7  0-7  2-4-  7-1 

Pb  6-1  1-8  0-8  3-2-  8-0 

All  measurements  are  expressed  as  a  percentage  of  the  standard  length.  The 
mouth  is  ventral,  the  anterior  margin  of  the  lower  jaw  is  curved  and  the  upper  and 
lower  lips  developed  but  not  lobed.  The  barbels  in  the  lectotype  show  signs  of 
damage.  The  head  is  pointed,  the  dorsal  margin  of  the  body  rises  in  a  smooth 
curve  from  the  snout  to  the  origin  of  the  dorsal  fin.  The  lateral  line  count  is  25  (f .9) 
or  26  (f.i)  (both  sides  of  the  fish  considered).  The  smallest  of  the  M.A.C.T.  speci- 
mens has  the  count  of  26  on  one  side.  There  are  4-5  scale  rows  between  the  dorsal 
mid-line  and  the  lateral  line  and  4-5  from  there  to  the  ventral  mid-line.  Two  and  a 
half  scales  are  between  the  lateral  line  and  the  base  of  the  pelvic  fin  and  12  scales 


K.    E.    BANISTER 


FIG.  90.     Barbus  ruasae  lectotype. 


encircle  the  caudal  peduncle.  The  striae  on  the  scales  are  parallel  and  much  wider 
apart  on  the  scales  of  the  pectoral  region  than  on  the  scales  of  the  caudal  peduncle. 
The  dorsal  fin  has  four  unbranched  rays,  the  last  is  ossified  into  a  strong  smooth 
spine  (x  =  21-0;  s.d.  =  2-9;  s.e.  =  1-3;  range  18-6-26-0).  There  are  8  (f.2), 
9  (f.2)  or  6  (f.i)  branched  rays.  The  dorsal  fin  with  six  branched  rays  showed 
obvious  signs  of  damage.  The  origin  of  the  dorsal  fin  is  above  or  slightly  anterior 
to  the  origin  of  the  pelvic  fin.  A  low  sheath  of  scales  surrounds  the  base  of  the 
dorsal  fin. 

The  anal  Jin  has  three  unfrranched  and  five  branched  rays. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  bones  are  moderately  slender,  the 
teeth  are  in  three  rows  of  5.3.2.  on  each  bone.  The  second  tooth  in  the  inner  row 
of  the  largest  M.A.C.T.  specimen  (No.  992214)  is  more  molariform  than  in  the 
lectotype. 

There  are  from  10  to  13  gill  rakers  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  In  alcohol  the  fish  present  a  silvery  appearance.  The  back  is  a  mid- 
brown  which  shades  to  silver  on  the  belly.  The  centres  of  the  scales  are  bright 
silvery  and  the  edges  are  darker.  Traces  of  dark  pigment  remain  in  the  middle  of 
the  caudal  fin,  the  distal  edge  of  the  dorsal  fin,  the  anal  fin  and  the  posterior  margins 
of  the  pectoral  and  pelvic  fins. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  ruasae  is  a  member  of  the  Barbus  intermedium 
complex  and  as  such  it  is  compared  with  and  distinguished  from  Barbus  acuticeps 
on  p.  8,  Barbus  altianalis  on  p.  27  and  Barbus  intermedium  on  p.  140. 

Pappenheim  &  Boulenger  (op.  cit.}  thought,  prophetically,  that  Barbus  ruasae 
showed  affinities  to  Barbus  leptosoma  (=  Barbus  intermedium). 

Very  low  lateral  line  counts  are  found  in  Barbus  mawambiensis.  This  species  is 
found  in  the  not-too-distant  Ituri  and  Ja  rivers  which  although  in  the  Congo  system 
do  not  flow  into  the  Lake  Victoria  basin.  I  do  not  have  enough  information  to 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


in 


determine  whether  or  not  there  is  any  relationship  between  Barbus  ruasae  and 
Barbus  mawambiensis. 

My  knowledge  of  the  Barbus  species  of  the  Congo  is  insufficient  to  even  hazard 
any  opinions  on  the  relationships  of  the  species  concerned. 


Barbus  somereni  Boulenger  1911 

Barbus  somereni  Blgr.,  1911,  Ann.  Mag.  nat.  Hist.  (8)  8  :  369  ;    Greenwood,  1966,  The  Fishes  of 

Uganda  2nd  ed.  :  69  ;    Banister,  1972,  Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  24  (5)  :  261-290. 
Barbus  altianalis  urundensis  David,  1937,  Revue  Zool.  Bot.  afr.  9  (4)  :  414. 
Barbus  urundensis  Poll,  1946,  Annls.  Mus.  r.  Congo  Beige  (i)  4  (3)  :  185-188. 

HOLOTYPE.     A  fish  of  172  mm  S.L.,  B.M.  (N.H.)  Reg.  No.  1911.7.26  :  i  from  the 
Sibwe  river,  Ruwenzori  mountains,  Uganda. 

DESCRIPTION.     The  description  is  based  on  51  specimens,  66-279  mm  S.L.     All 
measurements  are  expressed  as  percentages  of  the  standard  length. 


D 
H 

T 

10 

MW 

Pet 

CP1 

CPd 

Snt 

Ab 

Pb 


x 

29-5 

29-9 

7-6 

8-5 

7'5 

22-2 


II-6 

8-6 
7-8 
8-6 


s.d. 

2-9 

'4 

•4 

•i 

0-8 

•7 

•3 

0-7 

0-6 

i-o 

1-2 


s.e. 
0-4 

O-2 
O-2 
0-1 
O-I 
0-2 
O-2 
O-I 
O-I 
O-I 
0-2 


range 

25-4-38-4 

21-6-28-8 

5-2-10-6 

6-7-11-7 

6-3-  9-2 

18-4-28-0 

13-9-19-0 

10-4-13-2 

7-6-10-2 

5-6-  9-6 

6-3-11-6 


The  body  is  slightly  compressed  and  becomes  relatively  deeper  in  larger  fishes. 
This  and  the  relative  decrease  in  the  diameter  of  the  eye  with  an  increase  in  the 


FIG.  91.     Barbus  somereni  holotype  (from  Boulenger  1916). 


K.    E.    BANISTER 


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A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  113 

standard  length  are  the  only  obvious  signs  of  allometric  growth.  In  the  other 
measurements,  the  scatter  of  the  points  precludes  the  determination  of  the  direction 
of  the  allometry. 

The  mouth  is  sub-inferior  and  usually  has  continuous,  moderately  developed  lips, 
but  a  'rubber-lipped'  specimen  was  collected  in  the  Sibwe  river. 

Three  fish  from  the  Malagarasi  river  were  described  by  David  (1937)  as  Barbus 
altianalis  urundensis.  I  find  that  her  three  syntypes  (M.A.C.T.  Reg.  Nos.  46963- 
46965)  have  IV-8  (f.i),  IV-g  (f.2)  rays  in  the  dorsal  fin  (not  III-9  as  described) 
and  have  31  (f.i),  32  (f.i),  33  (f.i)  scales  in  the  lateral  line.  Her  counts  were  of 
the  total  number  of  scales. 

Squamation.     The  number  of  scales  in  the  lateral  line  ranges  from  26  to  34  : 

26  (f.i),  27  (f.2),  28  (13),  29  (f.5),  30  (f.n),  3i  (f-i2),  32  (f.7)>  33  (f-7)>  34  (f.i). 

The  pattern  of  striae  on  the  scales  varies  between  that  which  is  regarded  as  typi- 
cally radiate  and  that  which  is  typically  parallel.  There  is  a  general  tendency  for 
the  shoulder  scales  in  this  species  to  be  of  the  latter  type  and  the  belly  scales  to  be 
of  the  former.  The  striae  are,  however,  more  numerous  than  is  usual  for  the 
classical  radiately  striated  Barbus  (see  fig.  93). 

There  are  5-5  (sometimes  4-5  rarely  6-5)  scales  between  the  dorsal  mid-line  and 
the  lateral  line  and  4-5-6-5  (rarely  7-5)  scales  between  the  lateral  line  and  the  ventral 
mid-line.  The  specimens  from  the  Sibwe  and  Tokwe  rivers  (Lake  George  affluents) 
have  3-5  scales  between  the  lateral  line  and  the  pelvic  fin,  whilst  those  from  Mahembe 
and  the  Mutamphu  river  (Kagera  system)  have  2-5  scales  (rarely  2  or  3). 

There  are  12  scales  round  the  caudal  peduncle. 

Dorsal  fin.  There  are  four  unbranched  rays,  the  last  is  ossified  with  a  smooth 
posterior  margin.  This  is  also  tnie  for  the  type  specimen  although  Boulenger  (ignb) 
describes  only  three  unbranched  rays.  The  length  of  the  last  unbranched  ray  varies 
from  8-7  to  21-2  per  cent  of  the  standard  length  (x  =  13-8).  There  are  nine  or  ten 
branched  rays  except  in  one  fish  where  there  are  eight. 

The  anal  fin  has  three  unbranched  and  five  branched  rays. 

Coloration.  The  ripe-running  males  in  the  Sibwe  river  have  a  deep  olive-brown 
back  which  changes  sharply  into  an  ochrous  yellow  colour  on  the  flanks  and  belly. 
The  dark  olive  on  the  back  is  continued  into  the  lower  lobe  of  the  caudal  fin.  The 
upper  caudal  lobe  and  the  dorsal  fin  are  pale  brown.  This  pattern  of  a  dark  back 
and  dark  lower  caudal  lobe  persists  in  fishes  which  are  sexually  inactive  and  is  also 
visible  in  the  great  majority  of  preserved  specimens. 

GUI  rakers.  There  are  8-n  gill  rakers  on  the  lower  limb  of  the  first  arch.  The 
rakers  are  bilobed  with  the  sharply  triangular  outer  lobe  the  larger. 

The  pharyngeal  bones  and  teeth.  The  teeth  are  slightly  hooked  with  no  significant 
enlargement  or  molarization  of  the  second  tooth  on  the  inner  row  (fig.  94).  The 
pharyngeal  tooth  formula  is  2. 3. 5. -5.3. 2. 

DISTRIBUTION.  Specimens  are  known  in  the  Ruwenzori  area  from  the  following 
rivers  :  Sibwe  river,  Mubuku  river,  Tokwe  river  and  Kirimia  river.  In  the  Sibwe 
and  Mubuku  rivers  which  flow  into  the  northern  end  of  Lake  George,  Barbus  somereni 


K.   E.   BANISTER 


1  mm 


B,C. 


FIG.  93.  Scales  from  the  shoulders  of  A)  Barbus  bynni,  B)  B.  somereni,  C)  B.  humilis  to 
compare  the  striae  on  the  B.  somereni  scales  with  the  typically  'parallel'  and  'radiate' 
striae  patterns  shown  by  the  other  two  species. 


lives  in  the  faster-flowing  reaches  before  the  rivers  meander  into  the  lake.  Green- 
wood (1966)  gives  5500  feet  as  the  upper  limit  of  Barbus  somereni  distribution. 
Barbus  somereni  is  not  common  in  the  Mubuku  river  (see  Banister  1972). 

Barbus  somereni  has  also  been  found  in  the  Nyawarongo  (at  Mahembe)  and 
Akianaru  (Rwanda)  rivers,  in  the  Mutamphu  river  (12  km  from  Astrida  on  the 
road  to  Shangugu),  in  the  Chirangobwe  river  (Lake  Kivu  basin),  Mwogo  river 
(Kagera  system),  upper  Malagarasi  river  (Burundi),  Kitenge  river  (Ruzizi)  and  from 
the  Nyamagana  and  Nyakagunda  rivers  (Burundi). 

DIAGNOSIS  AND  AFFINITIES.  The  relationship  between  Barbus  somereni  (and  its 
relative  Barbus  mirabilis)  and  the  other  African  Barbus  species  is  uncertain.  The 
body  shape,  the  last  simple  dorsal  fin  ray,  the  number  of  dorsal  fin  rays,  the  nature 
of  the  striae  on  the  scales  and  the  colour  pattern  are  all  points  of  difference  between 
Barbus  somereni  and  the  members  of  the  Barbus  bynni  and  Barbus  intermedius  com- 
plexes (especially  Barbus  altianalis),  which  are  the  closest  groups  geographically. 

Barbus  somereni  is  a  species  which  lives  at  higher  altitudes  than,  for  example, 
Barbus  altianalis  eduardianus  in  the  Lake  George- Ruwenzori  Mountain  region. 
This  could  suggest  that  Barbus  somereni  has  been  displaced  by  Barbus  altianalis 
and,  if  so,  then  Barbus  somereni  is  a  longer  established  resident  of  the  area.  This 
view  is  strengthened  by  the  presence  of  Barbus  mirabilis  on  the  other  side  of  the 
rift  valley. 

Barbus  somereni  is  possibly  related  to  Varicorhinus  ruwenzorii.  These  two 
species  live  in  the  same  area,  have  the  same  colour  pattern  and  it  is  suggested 
(Banister  1972)  that  they  hybridize. 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS 


FIG.  94.     The  dorsal  and  lateral  aspects  of  the  left  pharyngeal  bone  from  Barbus  somereni. 


8* 


FIG.  95.     A  map  of  the  distribution  of  Barbus  somereni. 


u6  K.   E.    BANISTER 

Barbus  stappersii  Boulenger  1915 

Barbus  stappersii  Blgr.,  1915,  Revue  zool.  afr.  4  (2)  :  165  ;   Blgr.,  1920,  Revue  zool.  afr.  8  (i)  :  13  ; 

Jackson,  1961,  Fishes  of  Northern  Rhodesia  :  57. 
Barbus  curtus  Blgr.,  1915,  Revue  zool.  afr.  4  (2)  :  165  ;    Blgr.,  1920,  Revue  zool.  afr.  8  (i)  :  14  ; 

Jackson,  1959,  Occ.  Pap.  natn.  Mus.  Sth.  Rhod.  No.  236  :  298  ;    Soulsby,  1960,  Nth.  Rhod.  J. 

4  (4)  :  329,  fig.  10. 
Barbus  oxycephalus  Blgr.,  1915,  Revue  zool.  afr.  4  (2)  :  165  ;  Blgr.,  1920,  Revue  zool.  afr.  8  (i)  :  15  ; 

Soulsby,  1960,  Nth.  Rhod.  J.  (4)  :  329,  fig.  n. 
Barbus  moeruensis  Pellegrin,  1922,  Revue  zool.  afr.  10  (3)  :  273  ;   Pellegrin,  1928,  Annls.  Mus.  r. 

Congo  Beige  (i)  3  :  42. 

NOTES  ON  THE  SYNONYMY.  Jackson  (1961)  synonymized  Barbus  curtus,  Barbus 
oxycephalus  and  Barbus  moeruensis  with  Barbus  trachypterus  Blgr.  1915-  These 
four  species  and  Barbus  stappersii  were  each  described  from  single  specimens,  all 
from  Lake  Mweru,  and  all  except  Barbus  trachypterus  are  large  fish,  respectively 
their  standard  lengths  are  233  mm,  264  mm,  594  mm,  128  mm  and  283  mm. 

These  nominal  species  fall  into  two  obvious  groups,  one  with  the  caudal  peduncle 
as  deep  as  it  is  long  and  the  other  with  the  caudal  peduncle  much  longer  than  deep. 
The  former  group  consists  of  the  types  of  Barbus  stappersii,  Barbus  oxycephalus, 
Barbus  curtus  and  Barbus  moeruensis  whilst  the  latter  group  contains  Barbus 
trachypterus.  The  caudal  peduncle  depth  is  the  most  trenchant  difference  between 
the  two  groups  although  there  are  others.  The  size  difference  has  had  no  effect 
upon  the  dimensions  of  the  caudal  peduncle,  a  specimen  of  112  mm  S.L.  referable 
to  Barbus  stappersii  has  a  caudal  peduncle  deeper  than  it  is  long. 

Barbus  trachypterus  is  considered  here  to  be  a  valid  species  and  is  described  on 
p.  119. 

HOLOTYPE.     A  fish  of  283  mm  S.L.,  M.A.C.T.  No.  14250,  from  Lake  Mweru. 

DESCRIPTION.  The  description  is  based  on  nine  specimens  ranging  from  103  to 
594  mm  S.L. 

x  s.d.  s.e.  range 

D  35-6  3-8  i-i  30-0-40-7 

H  25-5  -2  0-4  25-0-28-2 

I  6-6  -3  0-4  4-1-  8-7 

IO  9-8  -4  0-4  7'7-I2>3 

MW  7-6  -4  0-4  6-2-10-8 

Pet  24-0  -2  0-7  18-6-26-0 

CP1  15-1  -4  0-5  13-0-17-9 

CPd  15-4  -7  0-5  12-0-17-4 

Snt  9-1  0-8  0-2  8-0-10-8 

Ab  2-7  0-9  0-3  1-8-  3-9 

Pb  3-6  0-7  0-2  2-8-  4-5 

They  are  stocky,  deep  fish  with  moderately  compressed  bodies.  The  ventral 
profile  is  gently  convex  from  the  mouth  to  the  anal  fin  whilst  the  dorsal  profile 
ascends  steeply  towards  the  dorsal  fin.  The  mouth  is  terminal,  the  lips  moderately 
developed  and  with  a  median  lobe  on  the  lower  lip.  All  the  examined  specimens 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


117 


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09 

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K.   E.    BANISTER 


FIG.  97.     The  dorsal  and  lateral  aspects  of  the  left  pharyngeal  bone  of  the  holotype  of 

Barbus  stappersii. 


have  lips  conforming  to  this  pattern,  except  for  a  specimen  from  Kilwa  in  which  the 
lips  are  thinner. 

The  caudal  peduncle  is  as  deep  as  it  is  long. 

Dorsal  fin.  The  leading  edge  of  the  dorsal  fin  is  slightly  in  advance  of  the  origin 
of  the  pelvic  fin.  A  low  basal  sheath  of  scales  is  present.  The  anterior  edge  of  the 
dorsal  fin  is  high  and  curved  so  that  the  concave  dorsal  margin  is  positioned  almost 
vertically.  The  dorsal  spine  is  moderately  well  ossified,  the  measurements  of  the 
non-flexible  basal  part  in  the  nine  specimens  are  x  =  16-0,  s.d.  =  3-88,  s.e.  =  1-37 
and  the  range  is  9-1  to  21-8  per  cent.  There  are  9  (f.2)  or  10  (i.j]  branched  rays. 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays. 

Squamation.  The  scales  have  numerous  parallel  striae.  The  lateral  line  has 
between  23  and  28  scales  :  23  (f.i),  24  (f-3),  25  (f.2),  26  (f.i),  27  (f.i),  28  (f.i). 
There  are  4-5  scale  rows  between  the  dorsal  mid-line  and  the  lateral  line  and 
4-5  (f.8)  or  5-5  (f.i)  scales  between  the  lateral  line  and  the  ventral  mid-line.  Two 
and  a  half  scales  are  present  between  the  lateral  line  and  the  pelvic  fin  base.  There 
are  12  scales  around  the  caudal  peduncle. 

Pharyngeal  teeth  and  bones.  The  pharyngeal  tooth  formula  is  2. 3. 5. -5.3. 2.,  the 
pharyngeal  bone  (fig.  97)  is  moderately  slender.  The  crowns  of  the  inner  row  of 
teeth  are  curved .  There  is  little  enlargement  of  the  second  tooth  on  the  inner  row, 
the  first  tooth  on  that  row  is  small  with  a  slightly  spoon-shaped  crown  and  it  is 
angled  towards  the  second  tooth.  The  succeeding  teeth  in  that  row  become  higher, 
more  slender  and  more  recurved.  The  tip  of  the  fifth  tooth  is  hooked  forwards. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 

^   0 


119 


FIG.  98.     A  map  of  the  distribution  of  Barbus  stappersii. 

Gill  rakers.     There  are  13  curved  gill  rakers  on  the  lower  limb  of  the  first  gill  arch. 

Coloration.  The  colour  in  alcohol-preserved  specimens  is  uniformly  brown,  with 
the  centre  of  each  scale  a  richer,  deeper  brown  than  the  posterior  margin. 

DISTRIBUTION.  This  species  is  known  from  Kilwa,  on  Lake  Mweru,  and  from  the 
Lubumbashi  region. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  stappersii  is  a  species  easily  recognized  by 
its  short,  deep  caudal  peduncle,  low  number  of  scales  in  the  lateral  line  series  (23-28) 
and  by  the  high  anterior  edge  of  the  dorsal  fin. 

Its  affinities  are  uncertain.  In  some  respects  (short,  deep  caudal  peduncle  and 
high  dorsal  fin)  it  resembles  specimens  of  Barbus  marequensis  A.  Smith  from  the 
Zambezi  river  (the  forms  described  as  Barbus  victoriae  Blgr.,  Barbus  altidorsalis 
Blgr.,  Barbus  chilotes  Blgr.,  Barbus  codringtonii  Blgr.  and  Barbus  fairbairnii  Blgr.). 
A  low  watershed  is  the  only  barrier  between  the  Upper  Zambezi  and  the  Lake 
Mweru  basin.  Until  I  have  examined  more  specimens  of  Barbus  marequensis  and 
Barbus  stappersii  I  am  reluctant  to  comment  further  on  any  relationship  between 
these  two  species. 


Barbus  trachypterus  Boulenger  1915 

Barbus  trachypterus  Blgr.,  1915,  Revue  zool.  afr.  4  (2)  :  164  ;    Jackson  (partim)  1961,  Fishes  of 

Northern  Rhodesia  :  58. 
Varicorhinus  bredoi  Poll,  1948,  Bull.  Mus.  r.  Hist.  nat.  Belg.  24  (21)  :  9. 

HOLOTYPE.     A  fish  of  128  mm  S.L.,  M.A.C.T.  No.  11380,  from  the  hydrographical 
station,  Lake  Mweru. 


120  K.    E.    BANISTER 

DESCRIPTION.     The  description  is  based  on  17  specimens  from  54  to  239  mm  S.L. 

n  x  s.d.  s.e.  range 

L  54    -239  mm 

D  17  27-9  1-6  0-4  25-6-31-4 

H  17  25-4  2-5  0-6  21-8-28-8 

I  17  5-4  i-o  0-2                  3-8-   7-4 

IO  17  7-7  1-3  0-3                 6-0-11-4 

MW  17  6-4  0-7  0-2                  5-4-   7-6 

Pet  17  2I-O  1-4  0-3  18-9-22-6 

CP1  17  17-4  1-3  0-3  14-5-19-3 

CPd  17  10-7  0-9  0-2  9-1-11-8 

Snt  17  8-2  i-i  0-2  6-5-11-4 

Ab  16  3-5  i-o  0-2  2-4-  5-5 

Pb  16  4-9  1-4  0-3  2-6-  8-0 

All  measurements  are  expressed  as  percentages  of  the  standard  length. 

The  body  is  slightly  compressed  with  a  level  or  gently  convex  ventral  profile  and 
a  dorsal  profile  which  rises  evenly  from  the  snout  to  the  dorsal  fin  origin  (fig.  99). 
The  mouth  is  ventral  and  semi-circular.  The  upper  lip  is  visible  in  the  ventral  view 
as  a  thin,  fleshy  surround  to  the  mouth.  The  thin  rostral  flap  reaches  down  to  the 
level  of  the  mouth.  The  anterior  edge  of  the  lower  jaw  is  strongly  convex  in  outline. 
In  small  fishes  a  fleshy  lower  lip,  often  with  a  small  mental  lobe,  is  present,  but  in 
larger  fishes  the  tendency  is  for  the  lower  jaw  to  have  a  flat,  cutting  anterior  margin. 
One  specimen  (M.A.C.T.  No.  129097)  has  well-developed  'rubber  lips'. 

Tubercles  are  present  on  the  snout  and  cheeks  of  several  specimens.  The 
tubercles  are  comparatively  small,  but  widespread  and  tend  to  coalesce.  In  speci- 
mens M.A.C.T.  Nos.  165254-165256  they  are  present  on  the  snout,  cheeks,  oper- 
culum  and  anal  fin  rays.  They  are  also  present  on  the  anal  fin  rays  of  the  holotype. 

In  three  comparatively  fresh  specimens  (M.A.C.T.  Nos.  165254-165256),  the  peri- 
toneum is  black. 

Squamation.  The  scales  possess  numerous  parallel  or,  at  the  most,  slightly  con- 
verging striae.  There  are  from  27  to  31  scales  in  the  lateral  series  :  27  (1.4), 
28  (f-4),  29  (f-5),  30  (f.2),  31  (f.2).  Twelve  scales  are  invariably  present  around  the 
caudal  peduncle.  There  are  4-5  (f.i5)  or  5-5  (f.i)  scales  between  the  dorsal  mid-line 
and  the  lateral  line  and  5-5  (f.io)  or  4-5  (f.3)  scale  rows  between  the  lateral  line  and 
the  ventral  mid-line  except  in  one  specimen  (M.A.C.T.  No.  129100  of  standard  length 
168  mm)  where  the  scales  on  the  chest  are  conspicuously  reduced.  There  are  2-5  or 
3  scales  between  the  lateral  line  and  the  base  of  the  pelvic  fins. 

Dorsal  Jin.  Except  for  the  holotype  of  Varicorhinus  bredoi  the  dorsal  fin  has  four 
unbranched  rays.  This  specimen  now  has  only  two  unbranched  rays  although  Poll 
(1948)  records  three.  There  is  little  doubt  that  the  reduction  in  the  number  of 
simple  rays  in  this  specimen  is  the  result  of  physical  damage.  The  last  unbranched 
ray  is  ossified  into  a  smooth,  often  slightly  curved  spine  :  x  =  20-0;  s.d.  =  2-48; 
s.e.  =  0-6;  range  =  i6-i-25-o.  The  dorsal  fin  is  slightly  in  advance  of  the  pelvic 
fin  origin.  There  are  8  (f.3),  9  (f.i2)  or  10  (f.2)  branched  rays. 

The  anal  fin  has  three  unbranched  and  five  branched  rays. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


c/i 
E 

00 


o 

E 

'o 

a 


K.    E.  BANISTER 


FIG.  100.     The  dorsal  and  lateral  aspects  of  the  left  pharyngeal  bone  from  the  figured 

specimen  of  Barbus  trachypterus. 


Pharyngeal  bones  and  teeth.  The  pharyngeal  tooth  formula  is  2.3. 5. -5.3. 2.  The 
pharyngeal  teeth  are  small,  slender  and  crowded  (fig.  100).  The  second  tooth  of  the 
inner  row  has  a  tendency  to  become  molariform  ;  in  six  of  the  nine  specimens 
examined  the  crown  is  rounded  and  flattened  at  the  top.  The  pharyngeal  bone  is 
thick  and  stout,  but  less  so  than  in  Barbus  caudovittatus  specimens  of  the  same  size 
(cf.  figs.  22  and  23). 

Gill  rakers.  The  gill  rakers  number  between  n  and  13  on  the  lower  arm  of  the 
first  gill  arch. 

Coloration.  The  colour  pattern  is  invariably  different  from  that  of  Barbus 
caudovittatus  with  which  some  of  these  specimens  have  been  confused.  The  black 
upper  half  of  the  sides  are  mid-brown  with  darker  brown  bases  to  the  scales.  The 
lower  part  of  the  flanks,  chest  and  belly  are  pinkish-brown.  Dark  pigment  is 
present  on  the  proximal  half  of  the  membrane  of  the  dorsal,  anal,  pelvic  and  pectoral 
fins.  The  caudal  fin  has  a  uniform  mid-brown  colouring.  The  colour  notes  are 
based  on  alcohol-preserved  specimens  (three  of  which  are  comparatively  recent), 
but  no  difficulty  was  experienced  in  separating  this  species,  on  colour  alone,  from 
Barbus  caudovittatus.  In  the  latter  species  the  dark  pigment  is  found  on  the  distal 
half  of  the  pectoral,  pelvic  and  anal  fins.  The  colour  pattern  is  sometimes  bleached 
out  in  specimens  that  have  been  badly  preserved  or  have  been  preserved  for  a  long 
time. 

DISTRIBUTION.  The  specimens  examined  came  from  Lake  Mweru,  from  between 
Kolwezi  and  Jadotville  on  the  Lualaba,  Upper  Katanga,  from  Mwena,  a  tributary 
of  the  Lufira,  Upper  Katanga,  from  Gombela,  Upper  Katanga,  from  Kabiyashi  on 


A   REVISION   OF   THE   LARGE   AFRICAN   BARBUS 


123 


FIG.  101.     A  map  of  the  distribution  of  Barbus  trachypterus. 

the  Luanza,   Upper  Katanga,   from  Ngundeulu,  Elizabethville   (=  Lubumbashi), 
and  Kando,  near  Tenke,  Upper  Katanga. 

AFFINITIES.  The  tubercles,  the  ventral  mouth  and  the  crowded  pharyngeal 
teeth  are  reminiscent  of  the  condition  found  in  many  species  of  the  genus  Varico- 
rhinus.  It  is  interesting  to  note  that  Poll  (1948)  thought  that  the  relationships  of 
Varicorhinus  bredoi  lay  with  Varicorhinus  stappersii  (here  considered  to  be  a  synonym 
of  Barbus  caudovittatus}  and  Varicorhinus  brucii  (which  was  considered  to  be  a 
variant  of  Barbus  marequensis  by  Groenewald  1958).  Barbus  trachypterus  specimens 
have  often  been  confused  with  Barbus  caudovittatus  specimens  but  can  be  dis- 
tinguished by  the  presence  of  a  stronger  dorsal  spine  and  by  a  different  colour 
pattern.  Both  Barbus  trachypterus  and  Barbus  caudovittatus  have  a  ventral,  curved 
mouth  and  possess  a  colour  pattern  unlike  many  of  the  east  African  species.  The 
affinities  of  Barbus  trachypterus  could  well  lie  with  Barbus  caudovittatus  but  much 
more  needs  to  be  known  about  the  Barbus  species  of  the  southern  and  eastern  parts 
of  the  Congo  before  a  more  informed  conclusion  can  be  drawn. 


Barbus  tropidolepis  Boulenger  1900 

Barbus  tropidolepis  Blgr.,  1900,  Annls.  Mus.  r.  Congo  Beige  Zool.  1  :  133,  pi.  xlix  ;    Poll,  1953, 

Result,  scient.  Explor.  hydrobiol.  Lac  Tanganyika  3  (5 A)  :  74. 
Varicorhinus  chapini  Nichols  &  LaMonte,  1950,  Proc.  biol.  Soc.  Wash.  63  :  175  (fide  Poll,  1952, 

Revue  Zool.  Bot.  afr.  (46)  3-4  :  222). 


I24 


K.   E.    BANISTER 


FIG.  102.     Barbus  tropidolepis  'Type'  specimen  (from  Boulenger  191  la). 


LECTOTYPE.  Boulenger  described  this  species  from  three  fish  from  Usambura, 
Lake  Tanganyika  [B.M.  (N.H.)  Reg.  Nos.  1906.9.6  :  19-21].  The  smallest  speci- 
men, 239  mm  S.L.,  is  recognized  as  the  lectotype. 

DESCRIPTION.  The  description  is  based  on  47  fish  of  99  to  365  mm  S.L.  The 
morphometric  data  are  expressed  in  tabular  form  below. 


D 

H 

I 

10 

MW 

Pet 

CP1 

CPd 

Snt 


28-8 
27-0 

9-4 
9-0 
6-4 

20-8 

14-9 

n-7 
8-4 


s.d. 

2-7 
1-4 


0-8 
i-o 

1-2 

0-9 

I-O 


s.e. 
0-6 


0-2 
O-2 


0-2 
O-2 


range 

25-0-33-7 

24-8-30-1 

7-0-11-0 

7-4-11-4 

5-2-  8-8 

19-1-23-9 

12-9-17-7 

10-1-13-7 

5-2-  9-3 


The  eye  is  large  and  frequently  protuberant.  The  characteristic  body  shape  is 
shown  in  fig.  102.  Although  the  anterior  barbel  is  invariably  absent  and  the 
posterior  represented  by  no  more  than  a  small  protrusion  (as  in  many  African 
Varicorhinus  species)  the  mouth  is  no  wider  than  in  most  Barbus.  Worthington  & 
Ricardo  (1937)  noted  that  the  degree  of  lip  development  varies  from  continuous  to 
discontinuous.  A  few  specimens  have  thickened  lips  but  in  none  of  the  fish  examined 
were  '  rubber  lips '  or  '  Varicorhinus-like '  lips  developed.  The  mouth  is  ventral 
under  an  obtuse  snout. 

Squamation.  The  lateral  line  has  from  39  to  44  scales  :  39  (f.2),  40  (f-9),  41  (f.n), 
42  (f.i-3),  43  (f.g),  44  (f-3).  There  are  8-5  (rarely  7-5)  scales  between  the  dorsal  mid- 
line  and  the  lateral  line  and  8-5  (rarely  7-5,  very  rarely  9-5)  between  the  lateral  line 


A   REVISION  OF  THE  LARGE  AFRICAN  BARBUS 


125 


FIG.  103.     The  dorsal  and  lateral  aspects  of  the  right  pharyngeal  bone  from 

Barbus  tropidolepis. 


and  the  ventral  mid-line.  Five  and  a  half  scale  rows  (rarely  4-5)  lie  between  the 
lateral  line  and  the  pelvic  fin  base.  Around  the  caudal  peduncle  are  16  (f-34), 
17  (f.6)  or  18  (1.7)  scales.  The  most  remarkable  feature  of  the  scales  of  Barbus 
tropidolepis  is  the  presence  of  longitudinal  folds  of  fat  which  form  well-marked 
ridges  on  the  body.  These  are  most  frequently  found  below  the  lateral  line  and 
from  the  middle  to  the  posterior  of  the  body.  Not  all  specimens  have  them,  e.g. 
B.M.  (N.H.)  Nos.  1936.6.15  :  596-598,  fishes  of  156  to  200  mm  S.L.  lack  the  ridges 
whilst  they  are  present  in  other  fish  of  less  than  100  mm  S.L. 

Dorsal  fin.  It  has  four  unbranched  rays  [not  three  as  reported  by  Boulenger 
(191  la)  and  Worthington  &  Ricardo  (1937)].  The  last  unbranched  ray  is  ossified 
to  form  a  smooth  stout  spine  which  varies  in  length  from  20-8  to  30  per  cent 
(x  =  25-8,  s.d.  =  2-7,  s.e.  =  0-6).  There  are  nine  (rarely  10)  branched  rays.  The 
dorsal  margin  of  the  fin  is  frequently  markedly  concave  with  its  posterior  corner 
extended  a  little  (see  fig.  102).  The  dorsal  fin  origin  is  in  advance  of  or  above  the 
origin  of  the  pelvic  fin. 

The  anal  fin  has  three  unbranched  rays  and  five  branched  rays. 

Pharyngeal  bones  and  teeth.  The  pharyngeal  teeth  are  molariform  (fig.  103).  The 
first  tooth  in  the  inner  row  is  veiy  small  and  in  a  few  cases  is  absent  although  a 
small  pinnacle  of  bone  marks  its  site.  The  second  tooth  is  large  with  a  slightly 


126 


K.    E.    BANISTER 


5  mm 

FIG.  104.     The  pharyngeal  bone  of  Barbus  tropidolepis  positioned  to  show  the  alignment 

of  the  tooth  rows. 


concave  crown.  The  third  and  fourth  teeth  are  wider  than  they  are  long  with 
concave  posterior  faces.  The  fifth  tooth  is  conical  and  may  be  recurved.  The 
alignment  of  the  second  and  third  rows  is  rather  unusual  in  that  the  first  tooth  of 
the  second  row  is  slightly  displaced  dorsally  so  that  both  rows  seem  to  radiate  from 
that  tooth.  In  the  other  species  under  consideration  the  second  and  third  rows 
are  distinct  and  parallel  (cf.  figs.  103  and  104).  The  pharyngeal  teeth  resemble 
those  of  Barbus  platyrhinus  (see-  p.  108) . 

Coloration.  Brown  or  olive-brown  above,  lighter  brown  or  silvery  below  in 
preserved  fishes. 

DISTRIBUTION.  This  species  is  endemic  to  the  Lake  Tanganyika  basin.  Poll 
(I953)  gives  a  full  list  of  the  localities  in  this  region  where  Barbus  tropidolepis  has 
been  captured. 

DIAGNOSIS  AND  AFFINITIES.  Barbus  tropidolepis  appears  to  occupy  a  rather 
isolated  position  among  East  African  Barbus  ;  only  Barbus  platyrhinus  has  some 
features  in  common.  The  large  number  of  scales,  the  development  of  ridges  of  fat 
on  the  scales  and  the  shape  of  the  pharyngeal  teeth  are  indicative  of  the  dissimi- 
larities between  this  species  and  those  of  the  neighbouring  areas  and  it  is  clearly 
distinct  from  Barbus  caudovittatus,  the  other  large  Barbus  species  in  the  lake. 

It  has  certain  features,  in  common  with  Varicorhinus  tanganicae,  viz.  a  high 
number  of  lateral  line  scales,  a  thick  dorsal  spine,  large  eyes  and  a  similar  snout. 
The  phyletic  significance  of  these  similarities  is  doubtful.  The  number  of  pharyngeal 
teeth  is  the  same  but  the  alignment  is  different. 

On  the  other  hand,  the  unique  specimen  of  Varicorhinus  chapini  is  definitely  a 
small  Barbus  tropidolepis  with  a  slightly  unusual  mouth.  The  measurements  of  this 
small  fish  do  not  differ  from  equal-sized  Barbus  tropidolepis  specimens. 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  127 

DISCUSSION 

Particularly  noteworthy  is  the  remarkable  degree  of  variation  in  lip  form,  body 
shape  and  dorsal  spine  strength  within  some  of  the  species  described  above  (e.g. 
Barbus  altianalis  and  Barbus  intermedius}. 

These  two  species  were  represented  by  very  large  series  of  specimens  (213  and  454 
fishes  respectively),  so  whether  the  same  degree  of  variability  would  be  shown  by 
other  species  when  equally  large  samples  were  studied  or  whether  these  two  species 
are  inherently  more  variable  than  the  other  described  species  cannot  at  the  moment 
be  determined.  Barbus  bynni  (59  specimens)  shows  much  less  variation.  The  same 
is  true  of  Barbus  oxyrhynchus,  which  although  more  variable  in  body  form  than 
Barbus  bynni  (cf.  the  type  of  Barbus  labiatus,  fig.  78,  with  the  type  of  Barbus  hindii, 
fig.  79)  does  not  approach  the  degree  of  variability  shown  by  Barbus  intermedius. 
Barbus  oxyrhynchus  is  represented  by  108  specimens,  i.e.  more  than  Barbus  bynni 
and  less  than  Barbus  altianalis  of  Barbus  intermedius.  Without  large  series  of 
specimens  this  matter  cannot  be  taken  further. 

The  intraspecific  variation  of  the  pharyngeal  teeth  is  as  large  as  the  interspecific 
variation  (excluding  Barbus  tropidolepis  and  Barbus  platyrhinus,  both  of  which  have 
a  very  large  degree  of  molarization  of  the  pharyngeal  teeth).  It  seems  that  the 
molarization  of,  particularly,  the  second  tooth  of  the  inner  row  is  of  no  significance 
in  establishing  the  identity  of  a  specimen.  The  range  in  pharyngeal  teeth  shapes 
found  in  Barbus  intermedius  (figs.  56,  57  and  58)  and  Barbus  altianalis  (figs.  13  and 
14)  show  this  well.  The  seemingly  random  occurrence  of  a  molariform  second  tooth 
in  the  inner  row  is  shown  by  the  series  of  pharyngeal  bones  of  two  of  the  subspecies 
of  Barbus  altianalis  (figs.  13  and  14,  also  p.  20) . 

It  was  suggested  above  (p.  16)  that  the  shape  and  strength  of  the  pharyngeal 
bones  and  teeth  might  be  influenced  by  the  diet.  The  pharyngeal  bones  of  Barbus 
altianalis  radcliffii  are  stouter  than  those  of  Barbus  altianalis  eduardianus.  Whether 
or  not  this  is  an  effect  of  dietary  differences  could  be  checked  comparatively  easily 
by  field  studies. 

It  is  known  that  water-snails  are  much  less  abundant  in  Lakes  Edward  and 
George  than  in  Lake  Victoria,  and  the  differences  between  the  pharyngeal  teeth  and 
bones  of  the  populations  of  the  cichlid  Astatoreochromis  alluaudi  in  these  lakes  has 
been  associated  with  this  fact.  Greenwood  (1964)  has  shown  the  differences  in  the 
stoutness  of  the  pharyngeal  teeth  and  bones  to  be  phenotypic.  Comparative  data 
on  the  diets  of  the  populations  of  Barbus  altianalis  in  these  lakes  are  lacking,  but 
the  striking  similarity  between  the  two  phenomena  is  suggestive. 

The  presence  of  massive  pharyngeal  bones  in  large,  deep-bodied  specimens 
(p.  16)  is  possibly  directly  related  to  body  depth.  The  ' surkis'  form  of  Barbus 
intermedius  has  been  shown  to  have  been  feeding  on  gastropod  molluscs  and  the 
likelihood  of  the  '  obesus '  form  of  Barbus  altianalis  radcliffii  also  having  gastropods 
as  an  important  part  of  its  diet  has  been  mentioned  above,  although  no  identifiable 
remains  were  found  in  its  alimentary  tract.  As  only  the  deep-bodied  examples  of 
populations  which  consume  gastropods  display  this  phenomenon,  it  is  more  likely 
that  the  length  of  the  pharyngeal  bone  is  associated  with  the  depth  of  the  body  as 
was  noted  on  p.  16. 


128  K.    E.    BANISTER 

Wunder  (1939)  experimented  on  Common  Carp  (Cyprinus  carpio  L.)  and  observed 
that  controlled  feeding  could  produce  a  'hunger'  form  and  a  'fattened'  form.  The 
former  resemble  the  'gorguarii'  form  of  Barbus  intermedium,  with  a  shallow  body  and 
a  relatively  long  head.  The  fattened  form  resembled  the  ' surkis'  form  of  Barbus 
intermedium  (fig.  32)  or  the  '  obesus '  form  of  Barbus  altianalis  (fig.  3)  in  having  a  deep 
body,  a  relatively  short  head  and  stubby  fins. 

It  is  not  possible  to  say  whether  greater  body  depth  in  a  fish  induces  longer  (but 
presumably  not  necessarily  stouter)  pharyngeal  bones  or  whether  extra  food  pro- 
duces a  deep-bodied  fish  in  which  the  pharyngeal  bone  size  relates  to  the  kind  and 
amount  of  food  rather  than  to  body  depth.  It  has  not  yet  been  possible  to  determine 
the  effect  that  a  deeper  body  has  on  the  depth  of  the  head.  It  would  be  interesting 
to  know  if  the  'gorguarii'  and  'surkis'  forms  of  Barbus  intermedium  represent  poorly 
fed  and  well-fed  populations.  This  could  possibly  be  answered  by  field  studies,  but 
until  that  time  one  can  do  little  but  accept  the  various  forms  as  different  phenotypes 
or  eco-phenotypes  of  variable  species.  It  is  important  to  note  that  the  deep-bodied 
forms  have  so  far  only  been  found  in  lakes  whilst  most  of  the  other  body  and  lip 
forms  occur  throughout  the  range  of  the  species.  There  is,  regrettably,  insufficient 
information  available  to  comment  further  on  the  variation  of  these  Barbus  species. 

As  in  the  small  Barbus  with  radiately  striated  scales  described  by  Greenwood 
(1962),  supra-specific  complexes  are  discernible  in  the  large  Barbus  described  above. 
These  complexes  (see  below)  are  difficult  to  define  precisely  since  they  are  based  on 
the  rather  amorphous  (although  useful)  concept  of  general  appearance.  It  must 
also  be  borne  in  mind  that  the  species  described  here  represent  only  a  fraction  of  the 
total  number  of  species  of  the  large  African  Barbus.  The  extent  and  composition 
of  the  supra-specific  assemblages  will  doubtless  have  to  be  re-evaluated  when  the 
phyletic  interrelationships  of  the  African  Barbus  are  better  understood. 

Two  complexes  can  be  recognized  amongst  the  species  studied  : 

1)  The  Barbus  intermedium  complex.     This  comprises  Barbus  intermedium,  Barbus 
altianalis,  Barbus  acuticeps  and  Barbus  ruasae  which  inhabit  most  of  the  rivers  of 
southern  and  eastern  Ethiopia  and  northern  Kenya,  the  Blue  Nile  system  (including 
Lakes  Rudolf  and  Baringo),  the  Lake  Victoria  basin  and  the  lakes  in  the  adjacent 
parts  of  the  western  rift  valley.     The  lateral  line  scale  counts  range  from  25  to  36 
but  are  most  frequently  28  to  32.     Typically,  the  body  is  shallow,  the  caudal 
peduncle  substantially  longer  than  deep  and  the  dorsal  spine  is  smooth,  strongly 
ossified  and  shorter  than  in  the  Barbus  bynni  complex  (see  below).     The  scales 
possess  many  parallel  or  slightly  converging  striae.     At  least  two  of  the  component 
species  (Barbus  intermedium  and  Barbus  altianalis)  are  noteworthy  for  their  high 
phenotypic  variability.     The  species  in  this  group  show  strong  gross  morphological 
resemblances  to  the  Barbus  of  western  and  southern  Saudi  Arabia  (e.g.  Barbus 
arabicus  Trewavas,  1939  and  some  as  yet  undescribed  species  from  Aden)  as  well 
as  to  species  like  Barbus  batesii  Blgr.  1903  from  South  Cameroons.     The  significance 
of  their  distribution  will  be  discussed  below. 

2)  The  Barbus  bynni  complex.     This  group  contains  Barbiis  bynni,  Barbus  ganan- 
ensis,  Barbus  oxyrhynchus  and  Barbus  longifilis  which  inhabit  the  White  and 
Albertine  Niles,  lakes  at  the  fringe  of  the  Blue  Nile  system  (Abaya  and  Rudolf), 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  129 

the  eastern  part  of  the  Juba  river,  the  Athi  and  Tana  rivers  and  the  Loama  and 
Kansehete  rivers  (upper  Congo  to  the  west  of  Lake  Kivu).  The  lateral  line  scale 
count  range  is  from  21  to  37,  the  most  southerly  species  (Barbus  oxyrhynchus) 
having  a  lower  range  than  the  rest,  modally  24-25  against  31-33.  The  trend 
towards  larger  scales  in  southerly  forms  has  been  noted  before  by  Greenwood  (1962) 
for  Barbus  paludinosus  and  Barbus  kerstenii.  The  scales  have  fewer  striae  than  in 
the  Barbus  intermedius  complex  and  the  striae  are  more  sinuous  The  caudal  peduncle 
is  deeper  in  relation  to  its  length  than  in  the  Barbus  intermedius  group  and  the  dorsal 
spine  is  noticeably  long  and  straight  (p.  28).  Barbus  oxyrhynchus  has  one  or  two 
more  branched  rays  in  the  dorsal  fin  than  the  other  species  in  this  group.  Barbus 
mariae  of  the  Athi  river  which  is  sympatric  with  Barbus  oxyrhynchus  does  not 
seem  to  have  much  in  common  with  the  bynni  complex,  except  a  long  dorsal  spine 
and  a  similar  lateral  line  scale  count  range. 

One  feature,  the  presence  of  small  scales,  is  common  to  several  lacustrine  species  : 
Barbus  tropidolepis,  Barbus  ethiopicus,  Barbus  platyrhinus  and  Barbus  microterolepis 
(see  below).  It  is  not  suggested  that  these  species  are  related,  merely  that  the 
character  has  been  acquired  independently  by  the  species  that  live  in,  and  probably 
evolved  in,  lakes.  Small-scaled,  large  Barbus  species  are  not  necessarily  lacustrine 
though,  as  some  of  the  fluviatile  South  African  species  (e.g.  Barbus  polylepis]  show. 
It  is  possible  that  the  modally  higher  lateral  line  scale  count  in  Barbus  altianalis 
amongst  the  Barbus  intermedius  complex  is  connected  with  its  confinement  in  lake 
basins.  However,  its  confinement  in  lake  basins  is  not  the  same  as  the  confinement 
in  lakes  of  Barbus  tropidolepis  etc.  There  are  populations  of  Barbus  altianalis 
radcliffii  which  appear  to  be  permanent  inhabitants  of  the  Kagera  river  (Dr  P.  H. 
Greenwood,  pers.  comm.).  Superficially  these  fishes  do  not  appear  to  differ  from 
those  in  Lake  Victoria,  but  no  detailed  investigations  have  been  carried  out.  The 
lacustrine  Barbus  altianalis  radcliffii  do  not  spend  all  their  time  in  the  lake,  they 
display  their  fluviatile  ancestry  by  ascending  rivers  to  breed  (Whitehead  1959). 

Barbus  tropidolepis,  it  should  be  noted,  also  ascends  rivers  to  breed  (Poll  1953) 
but  does  not  apparently  live  in  rivers  outside  of  the  breeding  season.  Nothing  is 
known  about  the  habits  of  Barbus  ethiopicus,  Barbus  platyrhinus,  nor  Barbus 
microterolepis. 

In  this  context  there  are  some  ideas  of  Farquharson  (1962)  that  must  be  con- 
sidered. He  discussed  the  distribution  in  South  Africa  of  the  small-scaled  Barbus 
species  (e.g.  Barbus  natalensis,  Barbus  holubi,  Barbus  capensis  and  Barbus  kimberley- 
ensis]  with  parallel  scale  striations.  From  their  widespread  distribution  and  endem- 
ism  he  suggested  that  an  ancient  dispersion  is  indicated.  If  this  is  so,  then  it  is 
possible  that  the  small-scaled  lacustrine  species  mentioned  above  are  relicts  of  this 
dispersion  southwards  (see  above).  I  have  not  been  able  to  compare  specimens  of 
the  small-scaled  South  African  species  with  the  small-scaled  species  from  Lakes 
Tanganyika  and  Zwai. 

Farquharson  (ibid.  :  247)  discusses  the  influence  of  water  temperature  on  scale 
number  and  lists  a  series  of  Barbus  species  arranged  in  order  of  localities  from  '  hot ' 
to  'cold'  showing  the  increase  in  the  number  of  lateral  line  scales.  Regrettably, 
the  situation  is  not  as  simple  as  this  as,  for  example,  the  review  by  Taning  (1952) 


130  K.    E.    BANISTER 

shows.  There  may  be  some  basic  truth  in  Farquharson's  idea  but  at  the  moment 
there  is  insufficient  data  to  put  forward  a  general  case. 

It  is  suggested  here  that  Barbus  altianalis  came  from  the  same  stock  as  Barbus 
intermedium  and  that  the  smaller  scales  developed  after  it  was  confined  to  the  lake 
basins,  subsequent  to  the  rift  valley  formation  (see  p.  22).  Barbus  microterolepis 
could  also  have  been  derived  from  Barbus  intermedium  in  this  fashion  (assuming  that 
it  is  a  good  species  and  not  a  hybrid  or  genetical  abnormality  -  see  p.  91). 

No  other  supra-specific  groups  are  recognizable  among  the  species  described.  The 
affinities  of  Barbus  stappersii  would  seem  to  be  with  the  Zambezi  species  Barbus 
marequensis.  Barbus  caudovittatus  is  probably  associated  with  Congo  species  and 
hence  outside  the  scope  of  this  paper.  Too  few  specimens  of  some  of  the  other 
species  are  available  to  be  able  to  evaluate  their  characters,  but  a  revision  of  the 
Congo  Barbus  may  give  some  indications  of  their  relationships. 

The  problematical  relationships  of  Barbus  macrolepis  may  not  be  solved  until  the 
Barbus  species  of  West  Africa  are  revised.  The  remarkable  similarity  of  this  species 
to  Barbus  lagensis  from  Nigeria  is  noted  above  (p.  83). 

Barbus  somereni  and  Barbus  mirabilis  do  not  seem  to  be  related  to  any  of  the  other 
species  described  in  this  paper.  Again,  a  study  of  the  Congo  fauna  may  reveal 
species  related  to  these  two. 

It  is,  perhaps,  interesting  to  compare  the  distribution  of  the  small  Barbus  species 
with  radiately  striated  scales  with  that  of  the  large  Barbus  species  described  above. 
The  small  Barbus  species  are  widely  distributed  and  several  species  are  found  in 
widely  separated  river  systems.  By  contrast  the  most  widespread  of  the  large 
Barbus  species  described  in  this  paper  are  found  in  palaeogeographically  closely 
related  and/or  adjacent  river  systems.  Presumably  the  small  Barbus  species  are 
able  to  travel  from  one  river  system  to  another  via  the  swamps  which  frequently 
form  the  watersheds.  Bell-Cross  (1965)  noted  that  the  only  Barbus  species  living 
in  the  watershed  on  the  Muhinga  plain  which  separates  the  Congo  from  the  Zambezi 
system  are  small,  'radiately  striated'  species.  The  large  Barbus  species  would 
seem  to  be  less  successful  at  negotiating  the  small  streams,  seasonal  ponds  and 
swamps  at  the  headwaters  of  the  river  systems.  Their  larger  size  must  be  a  con- 
tributing factor,  certainly  none  have  been  found  at  the  extremities  of  river  systems. 
Bell-Cross  (1960)  suggested  that  the  reason  why  some  fishes  did  not  cross  watersheds 
was  not  necessarily  because  of  their  physical  inability  to  do  so  but  because  of  some 
other  factor  like  behavioural  inhibitions.  Regrettably  there  have  been  very  few 
observations  on  this  problem.  Darlington  (1957  :  78-80),  however,  argues  that 
given  enough  time  primary  fresh-water  fishes  (e.g.  Ostariophysi)  can  overcome 
almost  any  physical  barriers  limiting  their  range.  Perhaps  the  time  scale  involved 
is  not  large  enough  to  be  applicable  to  the  large  Barbus  species  in  this  context. 

If  the  habitats  of  the  large  Barbus  species  are  likely  to  be  isolated,  the  possibility 
exists  that  the  relationships  of  various  species  might  prove  useful  as  indicators  of 
the  relationship  of  river  systems  and  vice  versa. 

If  the  rate  of  phenotypic  change  is  less  than  the  time  taken  for  rivers  to  be  dis- 
sociated then  it  is  feasible  to  think  that  the  now-separated  populations  of  fishes 
would  be  detectable  as  members  of  the  same  taxon.  The  general  principle  of  this 


A   REVISION   OF  THE  LARGE   AFRICAN  BARBUS  131 

idea  is  borne  out  in  certain  instances,  e.g.  between  Barbus  altianalis  and  Barbus 
intermedius,  between  Barbus  somereni  and  Barbus  mirabilis  and  between  the  mem- 
bers of  the  Barbus  bynni  complex.  The  geographical  evidence  for  this  idea  in  the 
Barbus  bynni  complex  is  lacking  in  detail,  but  there  is  nothing  geographically  incon- 
sistent in  the  former  association  of  their  river  systems  (see  p.  22). 

Giinther  (1869),  writing  on  the  Nile  fishes  collected  by  Petherick,  noted  that  the 
upper  Nile  fauna  is  related  more  closely  to  that  of  the  Palestinian  and  West  African 
rivers  than  to  the  fauna  of  Lake  Nyasa  (Malawi)  and  the  Zambezi  river.  By  1880 
Giinther  had  expanded  the  horizons  of  the  affinities  of  the  African  fresh-water  fish 
fauna.  He  thought  that  as  some  families  had  more  representatives  in  India  than 
in  Africa  they  probably  originated  in  India  or  derived  from  an  Indian  stock.  He 
also  considered  the  fauna  of  Jordan  and  Syria  to  have  so  many  African  representa- 
tives that  he  included  them  in  the  African  region  as  well  as  in  his  Euro-Asiatic  region. 

Gregory  (1896)  reviewed  the  evidence  for  the  fauna!  similarities  between  the  Nile 
and  Jordan  rivers  and  concluded  that  the  Jordan  river  has  species  in  common  with 
the  upper,  but  not  with  the  lower  Nile.  Because  of  this,  he  conjectured  that  in  the 
past  a  river  flowed  from  Lakes  Albert  and  Victoria,  which  then  had  no  outlet  to  the 
Nile,  into  Lake  Rudolf  and  then  via  the  Omo  and  Hawash  rivers  across  the  Afar 
depression  into  the  Red  Sea.  The  Red  Sea  trough  would,  he  thought,  at  that  time 
have  been  filled  with  fresh  water  and  with  an  extension  of  the  Jordan  river  flowing 
into  it. 

Nichols  &  Griscom  (1917),  with  a  great  many  more  species  at  their  disposal  than 
were  available  to  Gregory,  divided  the  fish  fauna  of  Africa  into  six  regions  and  com- 
mented on  the  relationships  of  one  faunal  area  to  another.  It  is  probably  significant 
that  the  boundary  line  between  their  '  Nile  and  North  East  Africa '  region  and  their 
'East  African'  region  follows  closely  the  boundary  between  the  ranges  of  Barbus 
intermedius  and  Barbus  oxyrhynchus.  The  fish  of  North  West  Africa,  they  noted, 
have  affinities  with  the  European  fishes.  The  fauna  of  the  'Nile  and  North  East 
Africa'  region  had  affinities  both  with  the  West  tropical  Africa  region  (Nichols 
and  Griscom  postulate  extensive  contact  in  the  past  between  the  Nile  and  the 
Congo)  and  northern  or  Asiatic  forms.  The  fauna  (particularly  the  Cyprinidae) 
of  the  Nile  and  North  East  Africa  region  ' .  .  .  seem  invariably  to  have  entered  Africa 
from  the  north-east.  They  predominate  in  the  Nile  basin  and  East  Africa,  reaching 
South  Africa  but  are  much  more  poorly  represented  in  West  Africa. ' 

Menon  (1964)  in  his  revision  of  the  cyprinid  genus  Garra  decided  that  the  African 
Garra  species  came  into  Africa  in  a  series  of  waves,  each  succeeding  wave  dispersing 
the  forms  that  came  in  the  preceding  wave.  His  postulated  routes  for  this  were 
either  via  the  Sinai  peninsula  or  via  Saudi  Arabia  and  Somaliland  before  these 
two  countries  were  separated  by  the  Red  Sea.  The  evidence  for  this  is  somewhat 
sparse,  the  idea  being  based  on  the  general  pattern  of  the  distribution  of  cyprinids 
in  Asia  and  the  pattern  of  distribution  of  fishes  in  Africa  with  the  most  archaic  forms 
in  the  west  (Darlington  1957  :  60). 

There  exist  in  south-western  Saudi  Arabia  and  Aden  some  Barbus  species  (Barbus 
arabicus  Trewavas  1939  -  and  some  undescribed  species)  which  would  fit  in  well 
with  the  Barbus  intermedius  supra-specific  complex.  The  question  that  cannot  yet 


132  K.   E.    BANISTER 

be  answered  is  whether  the  Arabian  species  represent  relict  populations  of  the 
migration  of  the  Barbus  intermedius  ancestral  stock  into  Africa  or  whether  they 
represent  a  radiation  from  Ethiopia  into  Arabia.  In  either  case  the  increase  in  the 
salinity  of  the  Red  Sea  (possibly  during  the  Pliocene  -  see  below)  severed  contact 
between  the  two  groups. 

The  geological  history  of  the  Red  Sea  is  not  known  in  great  detail,  but  enough  is 
known  to  enable  a  brief  history  to  be  constructed.  Dubetret  (1970)  considers  the 
Red  Sea  cut  across  the  Precambrian  rocks  forming  the  African  and  Arabian  shield. 
Up  to  the  Miocene  it  seems  that  marine  incursions  along  the  incipient  Red  Sea 
trench  were  confined  to  the  northern  end  (north  of  Quasir  27 °5'  N)  where  upper 
Cretaceous  marine  deposits  are  found.  The  Neogene  history  is  better  known. 

Dubetret  (op.  cit.}  dates  the  Red  Sea  from  the  Miocene.  Foundering  from  the 
Lower  Miocene  resulted  in  the  Mediterranean  Sea  flowing  into  the  Gulf  of  Suez  to 
approximately  the  southern  end  of  the  Red  Sea.  A  land  barrier  existed  at  the 
south  separating  the  Mediterranean  water  from  the  Indian  Ocean  water.  During 
the  late  Miocene  and  Pliocene  an  upward  earth  movement  in  the  Gulf  of  Suez  area 
cut  off  the  Red  Sea  from  the  Mediterranean. 

In  the  Pliocene  the  southern  land  barrier  sank  and  allowed  Indian  Ocean  water 
to  flow  into  the  inland  sea.  The  northern  land  barrier  prevented  any  contact  with 
the  Mediterranean. 

Botros  (1971)  thinks  that  a  series  of  fresh-water  lakes  was  present  in  the  southern 
part  of  the  Red  Sea  trench  during  the  Oligocene  and  late  Eocene,  whilst  the  northern 
end  was  an  arm  of  the  Mediterranean  Sea.  He  is  also  of  the  opinion  that  it  was 
possible  for  the  land  bridge  across  the  southern  end  of  the  Red  Sea  to  have  become 
apparent  again  during  the  Pleistocene  as  a  result  of  the  lowering  of  the  sea  level 
during  the  Ice  Ages. 

It  seems  then  that  what  is  'now  Arabia  (Roberts  1970  :  fig.  2)  did  not  separate 
from  the  horn  of  Africa  until  the  end  of  the  Miocene  or  early  Pliocene  arid  that  the 
two  land  masses  may  have  been  in  contact  again  during  the  Pleistocene.  It  can 
therefore  be  surmised  that  there  was  enough  contact  over  a  long  period  of  time  for 
Asiatic  cyprinids  to  have  migrated  into  Africa  or  vice  versa. 

The  pattern  of  distribution  of  the  supra-specific  complexes  defined  above  (fig. 
105)  refines  the  problem  but  does  not  solve  it.  The  geographically  most  compact 
supra-specific  complex  (the  Barbus  intermedius  complex)  is  in  northeast  Africa  (and  it 
is  suggested,  in  Saudi  Arabia).  This  is  ringed  to  the  west  and  south  by  the  rather 
more  fragmented  and  scattered  members  of  the  Barbus  bynni  supra-specific  complex. 
Still  more  scattered  are  the  localities  for  Barbus  somereni  and  Barbus  mirabilis. 
This  distribution  could  be  interpreted  as  the  result  of  a  series  of  invasions  from  the 
northeast.  Barbus  lagensis  and  Barbus  macrolepis,  if  they  are  as  closely  related  as 
the  available  evidence  suggests,  could  represent  the  remnants  of  an  early  scattering 
of  species  caused  by  these  invasions  (see  p.  83). 

The  reasons  for  this  apparent  effect  are  unknown,  but  it  could  be  related  to  the 
formation  of  topographical  changes  in  the  Ethiopian  highlands.  The  tectonic 
movements  must  have  had  a  profound  effect  in  altering  the  courses  of  rivers,  linking 
some  and  separating  others.  This  could  have  happened  more  than  once  and  over  a 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS 


133 


FIG.  105.  A  map  of  the  distribution  of  A)  the  Barbus  intermedius  supraspecific  complex 
(hatched  area),  B)  the  B.  bynni  supraspecific  complex  (dark  area),  C)  B.  somereni  and 
B.  mirabilis  (solid  circles). 

9 


134  K-   E-   BANISTER 

long  period  of  time,  which  could  ultimately  cause  speciation  within  a  previously 
uniform  population.  Unfortunately  the  geological  history  of  this  region  is  poorly 
known.  Migration  west  and  south  would  scatter  the  daughter  species.  Any  sub- 
sequent waves  of  immigration  in  this  region  might  hence  be  subject  to  the  same 
kinds  of  isolation  as  the  earlier  waves  and  further  supra-specific  complexes  would 
result. 

There  is  evidence  to  suggest  that  fishes  have  been  able  to  move  from  northeast 
Africa  into  the  middle  east.  The  presence  of  Tilapia  species  in  Jordan  is  an  example 
of  this.  The  only  conclusions  that  can  be  drawn  about  the  large  Barbus  species  is 
that  they  were  at  some  stage  able  to  move  between  the  horn  of  Africa  and  Saudi 
Arabia.  It  would  need  fossil  evidence  to  show  in  which  direction  the  movement 
took  place.  Brown  (1970)  reports  the  presence  of  Barbus  species  and  Tilapia 
species  in  the  late  Oligocene  or  Miocene  of  Jizan  (Saudi  Arabia,  just  north  of  the 
Yemen  border). 

The  large  Barbus  species  of  Africa  are,  as  a  group,  quite  distinct  from  the  majority 
of  species  living  in  the  Tigris  and  Euphrates  rivers  (the  nearest  region  with  a  good 
cyprinid  fauna) .  The  Tigris  and  Euphrates  species  mostly  resemble  Barbus  barbus  L. 
of  Europe  in  possessing  relatively  small  scales  and  a  serrated  dorsal  spine.  Of  the 
smooth  dorsal  spined  species  from  Syria,  Barbus  canis  C.  &  V.  (Tor  canis  of  Karaman 
1971)  most  closely  resembles  the  African  species,  especially  the  Barbus  intermedium 
complex.  An  isolated  population  of  Barbus  canis  from  Khamis  Mushyat  (i8°i7'  N, 
42°34'  E  -  about  400  km  southeast  of  Jeddah,  Saudi  Arabia)  is  the  population  closest 
to  the  African  continent.  The  members  of  this  population  are  less  like  the  African 
Barbus  species  than  are  the  Syrian  populations  as  they  lack  the  anterior  pair  of 
barbels.  The  fish  fauna  of  Saudi  Arabia  is  sparse  and  too  poorly  collected  for 
opinions  to  be  formed  on  whether  the  absence  of  the  anterior  barbels  in  the  Khamish 
Mushyat  population  is  a  local'  phenomenon  or  part  of  a  clinal  phenomenon.  The 
Barbus  species  of  Aden  are,  as  has  been  mentioned  above,  extremely  similar  to 
Barbus  intermedius. 

The  presence  of  Barbus  species  in  Saudi  Arabia  which  closely  resemble  some  of 
the  African  species  presents  two  possibilities  about  the  origin  of  the  African 
representatives. 

Firstly,  did  the  Barbus  species  now  typical  of  Africa  evolve  in  Africa  from  dis- 
similar ancestors  of  which  now  no  trace  remains?  If  so,  then  the  Arabian  and 
Syrian  smooth-spined  Barbus  species  could  represent  the  relicts  of  a  migration  from 
Africa.  The  centre  of  this  particular  radiation  could  well  have  been  in  Ethiopia  as 
the  African  and  Arabian  land  masses  must  have  been  in  contact.  Therefore  the 
status  of  the  Arabian  and  Syrian  species  would  be  the  same  as  Barbus  altianalis  and 
Barbus  ruandae,  i.e.  the  members  of  the  Barbus  intermedius  supra-specific  complex. 

The  other  possibility  is  that  the  facies  characteristic  of  the  African  Barbus  species 
were  to  some  extent  present  in  the  ancestral  stock.  If  this  were  so,  and  if  the 
ancestral  stock  was  not  originally  African,  could  the  Arabian  and  Syrian  species  of 
the  Barbus  canis  type  represent  remnants  of  the  ancestral  stock? 

Not  enough  is  known  at  the  moment  to  be  able  to  decide  which  of  these  two 
possibilities  (if  either)  represents  the  truth  and  it  is  quite  possible  that  the  notion  of 


A   REVISION   OF  THE   LARGE   AFRICAN  BARBUS  135 

a  series  of  migrations  all  in  the  same  direction  is  far  too  simple.  The  ideas  in  the 
previous  two  paragraphs  refer  only  to  the  putative  last  migration  that  produced  the 
species  forming  the  Barbus  intermedium  supra-specific  complex.  Of  the  even  earlier 
migrations  and  scatterings  conjectured  from  the  distribution  of  the  extant  species 
there  is  no  trace,  but  this  does  not  affect  the  equally  likely  (on  available  evidence) 
alternative  ideas  on  the  relationship  of  the  Barbus  intermedium  complex  to  the 
relevant  Barbus  species  of  Saudi  Arabia  and  Syria. 

However,  no  firm  conclusions  can  be  drawn  until  the  fossil  record  and  the  phylo- 
geny  of  this  group  of  Barbus  species  are  better  known.  For  a  full  understanding 
of  the  phylogeny  and  distribution  of  the  African  Barbus  species,  the  relationship  of 
the  small  'radiately  striated'  Barbus  species  to  the  large  Barbus  species  must  be 
determined  as  well  as  the  relationships  of  the  African  Barbus  species  sensu  lato  to 
the  Barbus  species  of  Europe  and  the  Near  East.  The  distributional  problems 
relate  principally  to  the  role  of  the  closely  related  cyprinid  genera,  i.e.  why  should 
there  be  a  relative  paucity  of  large  Barbus  species  in  west  Africa  but  a  comparative 
abundance  of  Varicorhinus  species  and  Labeo  species?  The  reverse  situation  exists 
in  the  east.  In  all  probability  the  solutions  to  these  problems  lie  as  much  in  the 
sphere  of  the  field  worker  as  in  the  sphere  of  the  museum  worker. 

ACKNOWLEDGEMENTS 

Many  people  have  helped  me  in  the  production  of  this  paper  and  it  is  with  con- 
siderable pleasure  that  I  take  this  opportunity  of  thanking  them  all. 

My  senior  colleagues  Dr  P.  H.  Greenwood  and  Dr  E.  Trewavas  have  spent  many 
hours  discussing  speciation  and  zoogeography  and  have  generously  given  me  the 
benefit  of  their  knowledge  and  experience  of  African  fish.  Dr  M.  Poll  of  the  Central 
African  Museum,  Tervuren,  Belgium,  gave  me  facilities  to  study  the  collections  in 
his  charge  as  well  as  readily  sending  specimens.  Dr  W.  Klausewitz  of  the  Sencken- 
berg  Museum,  Frankfurt,  Dr  E.  Tortonese  of  Genoa,  Dr  C.  Karrer  of  Berlin  and 
Dr  J.  C.  Tyler  of  Philadelphia  respectively  allowed  me  to  examine  the  collections 
of  Riippell,  Vinciguerra,  Pappenheim  and  Donaldson-Smith.  Through  the  good 
offices  of  Dr  Letitzia  Ferrara  of  the  Stazione  Idrobiologia,  Rome,  I  was  able  to 
examine  Dr  G.  Bini's  extensive  collections  from  Ethiopia. 

I  want  to  express  my  gratitude  to  Miss  M.  A.  McLellan  for  the  help  she  gave  me 
in  preparing  the  graphs  and  with  the  calculations.  The  drawings  of  the  pharyngeal 
bones  are  the  work  of  Mr  G.  J.  Howes  and  the  uncredited  drawings  of  whole  fish  are 
the  work  of  Mrs  S.  Chambers.  Mr  J.  Chambers  prepared  some  alizarin  specimens. 

Dr  W.  W.  Bishop  has  helped  considerably  in  advising  me  and  giving  me  infor- 
mation on  stratigraphical  and  palaeogeographical  matters.  The  staff  of  the  Royal 
Geographical  Society  deserve  my  heartiest  thanks  in  tracing  localities  that  I  had 
thought  were  untraceable. 

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APPENDIX   i 

An  artificial  key  to  the  Barbus  species  described  in  this  paper 

Li  :  >  46 

Lake  Zwai  basin     .......  B.  ethiopicus 

Li  :  38-44 

a)  No  anterior  barbel ;    posterior  barbel  minute  ;    ridges  of  fat  on  scales  of 

adults 

L.  Tanganyika  basin        .          .          .          .          .  B.  tropidolepis 

b)  Anterior  barbel  present 

i)   IO  more  than  8  per  cent  S.L.  ;   heavy-bodied,  bulky  appearance  ; 
pharyngeal  bones  short  and  thick,  pharyngeal  teeth  molariform 

L.  Tanganyika  basin        .          .          .          .          .  B.  platyrhinus 

ii)    IO  less  than  8  per  cent  S.L.  ;  compressed,  slender  body  ;  pharyngeal 
teeth  thin  and  hooked 

L.  Zwai  basin          ......  B.  microterolepis 

Li  :  <  38 

a)  Last    unbranched    dorsal    fin    ray    weak    and    flexible    with    persistent 

articulations 

i)    CP1  >  CPd  ;    body  shallow  ;    L24~3O  (most  often  26-28),  striae  on 
scales  parallel 

L.  Tanganyika  basin  and  upper  reaches  of  Congo        .      B.  caudovittatus 
ii)    CP1  —  CPd  ;   body  deep  ;   scales  with  few  converging  striae  (fig.  64) 

Lower  Malagarasi  and  Tanzanian  coastal  rivers  .          B.  macrolepis 

b)  Last  unbranched  dorsal  fin  ray  ossified  without  articulations,  at  least  in 

basal  half 
i)   Body  typically  deep  ;   CP1  -  CPd 

t  Dorsal  spine  thin,  only  ossified  basally 

L.  Mweru  region     .          .          .          .          .          .          .  B.  stappersii 

If  Dorsal  spine  long,  thick  and  strong      .          .          .  B.  bynni  complex 

(except  in  B.  pagenstecheri)  (see  below) 

ii)    Body  typically  shallow  ;   CP1  >  CPd 

t  Dorsal  spine  massive,  long  ;    upper  jaw  curves  down  to  cover  ft 

the  lower  jaw  laterally  giving  a  'clupeoid'  appearance  to 
the  mouth 
Athi-Tana  system  .          .          .          .          .          .  B.  mariae 

ft  Dorsal  spine  not  as  above 

%  mouth  ventral  %% 

D  straight  edge  to  square  LJ  ;  DD 

very  small  barbels 
Rwanda  .         B.  microbarbis 


140  K.    E.    BANISTER 

DD  mouth  semicircular 

*  Dark  pigment  of  distal  portions  of  pectoral,  pelvic 

and  anal  fins 
Upper  Congo,  W.  of  L.  Kivu   ....  B.  paucisquamatus 

**  Dark  pigment  on  proximal  parts  of  pectoral,  pelvic 

and  anal  fins 
L.  Mweru  region     .          .          .          .          .          .  B.  trachypterus 

JJ  mouth  not  as  above 

|  striae  on  scales  vary  with  position  of  scale  on  body,  Bfl] 

typically  striae  more  radiate  on   belly  than  on 
back  ;    DIV-g  or  10 
O  Bulky  body  (fig.  74) 
West  of  rift  valley  .          .          .          .          .          .  B.  mirabilis 

O  O  Body  not  bulky 

Ruwenzori  mountains  and  upper  Malagarasi       .          .  B.  somereni 

••  striae  ±  parallel  and  sinuous,  no  great  variation  in 
pattern  with  position  on  body  ;    DIV-8  or  9 

B.  intermedius 
complex  (see  below) 

Barbus  bynni  complex 

1)  DSP,  x  =  30-0,  spine  strong      ..........          2) 

Nile,  L.  Albert,  L.  Abaya,  L.  Rudolf          .  .  B.  bynni 

2)  DSp,  x  =  25-0-30-0,  strong       ..........          3) 

a)  Ab,  x  =  9  (range  8-10)  ;    Pb,  x  =  n  (range  9-12). 

Pet  21-24 

Congo  W.  of  L.  Kivu       .          .          .          .          .          .  B.  longifilis 

b)  Ab,  x  =  6  (range  3-10)  ;   Pb  x  =  7  (range  3-12). 

Pet  22-28,  x  =  24-4 

Athi  and  Tana  systems  .....       J5.  oxyrhynchus 

3)  DSp  25-0  or  less,  but  weaker  than  above 

Juba  river      ........          B.  gananensis 

4)  DSp  20-0  or  less,  shallow  body 

Kilimanjaro  region  .          .          .          .          .  B.  pagenstecheri 


Barbus  intermedius  complex 

1)  Dorsal  profile  of  head  concave  ;    mouth  opens  antero-dorsally  ;    nuchal  hump 

present 

Li  27-30  ;   DSp,  x  =  22-3 

Rwanda  and  Burundi      .          .          .          .          .          .  B.  acuticeps 

2)  Dorsal  profile  of  head  not  concave,  mouth  sub-terminal  or  terminal 

i)       Li  25-26  ;   pharyngeal  teeth  tend  to  be  molariform 

Rwanda          .........       B.  ruasae 

ii)       Li  28-35  (most  often  30-34,  m  =  32) 

DSp,  x  =  16-9  ;   Ab,  x  =  4-3  ;   Pb,  x  =  5-4 

Lakes  Victoria,  Kivu,  Edward  and  George          .          .  B.  altianalis 

iii)       Li  26-34  (most  often  28-32,  m  =  29) 

DSp,  x  =  20-1  ;   Ab,  x  =  5-8  ;   Pb,  x  =  6-9 

Blue  Nile,  Ethiopia,  Lakes  Rudolf  and  Baringo,  western  part 

of  Juba       .          .          .          .          .          .          .  B.  intermedius 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  141 

APPENDIX   2 

Gazetteer 

As  I  have  found  it  very  difficult  (and  in  some  cases  impossible)  to  find  some  of  the 
localities  mentioned  by  authors  I  have  compiled  this  gazetteer  giving  as  many 
details  as  I  can  find  of  the  collecting  sites.  Grid  references  are  given  where  possible. 
The  details  of  the  rivers  in  Ethiopia  come  entirely  from  Zaphiro's  notebook. 
Different  cultures  occupying  an  area  have  sometimes  changed  the  names  of  lakes 
or  rivers.  Where  possible  all  the  variants  are  given. 

Abaya  =  Abbaya,  the  northern  lake  of  a  pair  of  lakes.     The  name  has  been  loosely  applied  to 

either  or  both  lakes  in  the  southern  part  of  the  Ethiopian  rift  valley.     G.R.  6°3o'  N  3o°oo'  E. 

(See  Chiamo,  Ganjule,  Margharita,  Ruspoli.) 
Akaki,  a  river,  10-12  m  wide,  due  south  of  Addis  Ababa,  it  runs  from  Legadi  south  to  the 

Hawash. 
Alaba,  a  river,  some  20  m  wide,  runs  from  the  Kambata  plains  to  Lake  Ganjule.     Collecting 

altitude  6000  feet. 

Arba  minch,  tributary  of  Lake  Ganjule. 
Avakubi,  Ituri  system  (Zaire).     G.R.  i°24' N,  27°4o'  E. 
Awata,  a  tributary  of  the  Juba  river.     G.R.  6°O5'  N,  39°2o'  E. 
Barja,  a  river,  10  m  wide,  collecting  altitude  4250  feet.     It  runs  from  the  hills  of  Sangana  and 

Bako  to  the  Zuja  river  (Lake  Stephanie  basin). 
Basso  Ebor,  an  old  name  for  Lake  Stephanie. 
Basso  Narok,  an  old  name  for  Lake  Rudolf. 

Bis (s) an  Guarrica,  a  tributary  of  the  Sagan  river  to  the  south  of  Lake  Abaya. 
Bobandana,  Lake  Kivu  basin.     G.R.  i°38'  S,  29°oo'  E. 

Bushiame  river,  Sankuru  system.     G.R.  6°oo'  S,  24°5o'  E  to  8°oo'  S,  23°oo'  E. 
Burka,  an  affluent  of  Lake  Ganjule. 
Chiamo  (=  Ciamo),  Lake  Ganjule. 
Chirangobo,  an  affluent  of  Lake  Kivu. 

Didessa,  a  river,  some  60  m  wide,  that  runs  from  Guma  to  the  Blue  Nile. 
Elgo,  a  fast-flowing  river,  15  m  wide,  collecting  altitude  3000  feet,  it  flows  from  the  Gamu  hills 

eastwards  to  Lake  Abaya. 
Ergino,  a  river,  20  m  wide,  collecting  altitude  3000  feet,  flows  from  the  Basketo  hills  northwards 

to  the  Omo. 

Errer,  a  river  in  the  Webi  Shebeli  system.     G.R.  9°oo'  N,  42°2o'  E. 
G.  el  Narua,  Guasso  el  Narua.     G.R.  o°i6'  N,  36°i8'  E. 

G.  Nyuki,  Guasso  Nyuki,  mouth  at  o°28'  N,  36°o8'  E  (probably  a  swampy  locality). 
Ganda,  a  river,  15  m  wide,  collecting  altitude  3000  feet,  flows  from  the  hills  of  Gama  to  Lake 

Ganjule. 
Ganjule,  the  southern  lake  of  the  pair  of  which  the  northern  lake  is  Abaya  (also  called  Chiamo, 

Ruspoli).     G.R.  6°50'  N,  37°4o'  E. 
Gato,  a  fast  flowing  river,  15  m  wide,  which  rises  in  the  hills  of  Gandulla  and  flows  west  to  the 

Sagan.     Collecting  altitude  3700  feet.     (Lake  Stephanie  basin.) 
Gibe,  a  fast-flowing  stream,  20  m  wide,  which  flows  directly  south  from  the  plains  of  Gorombi 

(altitude  2800  feet)  (supposed  to  be  the  source  of  the  Omo). 
Gofa,  a  river,  Hawash  system. 
Gombela.     G.R.  io°5o'  S,  27°5o'  E. 
Gudar   (=  Gudr),    a   stream   flowing   north   from   the   Roggle  mountains   to   the   Blue   Nile. 

Collecting  altitude  3400  feet. 

Hawash,  most  collections  in  this  river  were  made  by  Zaphiro  at  Zeluka,  altitude  4100  feet. 
Jerrer,  exact  locality  untraceable.     Mount  Jerrer  is  32  km  southwest  of  Addis  Ababa,  according 

to  Zaphiro's  notebook,  the  river  is  therefore  apparently  in  the  Webi  Shebeli  system. 


142  K.    E.    BANISTER 

Juju,  a  river,  10  m  wide,  flowing  from  the  Guma  hills  to  the  Blue  Nile.     Collecting  altitude 

2000  feet. 

Kabiashyia,  on  the  Luanza  river,  an  affluent  to  the  northern  edge  of  Lake  Mweru. 
Kando,  near  Tenke.     G.R.  io°25'  S,  26°io'  E. 

Kansihiti,  a  river,  an  affluent  of  the  Luhoho  Congo,  due  west  of  Lake  Kivu. 
Kassam,  a  river,  Hawash  system.     G.R.  9°O5'  N,  39°35'  E. 

Kibonoto  ( =  Kibongoto),  an  affluent  of  the  Sanya  river,  Pangani  system.     G.R.  3°i  i'  S,  37°o6'  E. 
Kibwesi,  a  river  in  the  Athi  system.     G.R.  ±  2°25'  S,  37°56'  E. 
Kiyimbi,  a  river,  Loama-Lualaba  Congo.     G.R.  5°oo'  S,  28°59'  E. 
Koki,  a  river,  an  affluent  of  Lake  Tanganyika.     G.R.  6°O3'  S,  29°O5'  E. 
Laikipa  (Leikipa),  an  escarpment.     G.R.  o°25'  N,  36°io'  E. 
Loama,  an  affluent  of  the  Luhoho  Congo,  due  west  of  Lake  Kivu. 
Luembe,  a  river  =  Cashimo  river.     G.R.  8°oo'  S,  2i°35'  E. 
Lufiro,  a  river.     G.R.  2°4o'  S,  29°oo'  E. 
Luilu,  a  river,  Sankuru  system.     G.R.  7°3o'  S,  23°3o'  E. 
Lusine,  a  river,  Rwanda,  flows  from  Lake  Mohashi  into  the  Akangaru.     G.R.  2°3o'  S  to  6°oo'  S, 

30°45'  E. 
Mahembe,  on  the  Nyawarongo  river,  Kagera  system,  Lake  Victoria  basin.     G.R.  ±  i°52'  S, 

29°54'  E. 
Maki,  a   fast-flowing  stream   that  originates   in   the  hills   of  Goraza   and   flows   into   Lake 

Zwai. 

Makindu,  a  river.     G.R.  2°O9'  S,  37°35'  E. 

Malawa,  a  river,  Lake  Victoria  basin.     G.R.  o°4o'  N,  35°3o'  E. 
Margharita,  lake  =  Lake  Abaya. 
Metti,  a  fast-flowing  river,  20  m  wide,  altitude  3500  feet.     Flows  from  Tuludimtu  northwards 

to  the  Gudar  river.     (Blue  Nile.) 
Mkunga  (=  Mukungwa),  a  river,  near  Ruasa,  northwest  Rwanda,  runs  from  the  southeastern 

part  of  Lake  Luhondo  to  the  Nyawarongo.     G.R.  i°35'  S,  29°4o'  E. 
Modjo,  a  river,  Webi  Shebeli  system. 
Mogre,  a  river,  tributary  of  the  Blue  Nile. 
Mutamphu,  a  river,  a  tributary  of  the  River  Akangaru,  Kagera  system.     Specimens  with  this 

locality  were  collected  about   i 2  km  along  the  road  to  Shangugu  from  Butare.     G.R. 

±  2°43'  S,  29043'  E. 
Mwogo,  an  affluent  of  the  Kagera. 
Narok,  a  river.     G.R.  o°32'  N,  36°52'  E. 
Nyabarongo,  a  river  in  the  Luhoho  Congo  system. 
Nyabugogo,  an  affluent  of  the  Lusine  river. 
Nyawarongo,  Kagera  system. 
Nyundeulu.     G.R.  io°58'  S,  25°5o'  E. 
Rugwero.     Lake,  Rwanda.     G.R.  2°25'  S,  3O°2o'  E. 
Ruspoli  =  Lake  Ganjule. 
Sagan,  a  fast-flowing  stream,  15  m  wide,  at  Wondo  (altitude  2800  feet)  where  most  collecting 

was  done.     Blue  Nile  system. 

Sang6,  an  affluent  of  the  Ruzizi.     G.R.  3°O4'  S,  29°o8'  E. 
Sir 6  (=  Siri),  a  stream  flowing  from  the  Gamu  hills  eastwards  to  Lake  Abaya,  collecting  altitude 

3000  feet. 

Suksuki,  a  river  linking  Lake  Zwai  and  Lake  Suksuki. 

Tchitatu,  a  river,  near  Luachimo,  Upper  Kasai  region.     G.R.  7°4o'  S,  2O°5o'  E. 
Tokwe,  a  river,  near  Bwambe,  Uganda. 
Tsavo,  a  river.     G.R.  2°59'  S,  38°O2'  E. 

Tshikapa,  a  river,  Upper  Kasai  (Congo)  system.     G.R.  6°28'  S,  2o°48'  E. 

Uaso  narok,  a  river  flowing  from  the  Laikipa  escarpment  northeastwards  to  the  Uaso  Nyiro. 
Urgessa,  a  river,  15  m  wide,  flowing  northwest  to  the  Wama  river  (Blue  Nile  system). 
Wondinak,  a  small  stream  flowing  northwest  into  the  Gibe  river  (Omo  system). 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  143 

Zeissi,  a  turbulent  stream,  about  10  m  wide,  flowing  from  the  Zeissi  hills  eastwards  to  Lake 

Ganjule.     G.R.  i°52' N,  37^9' E. 
Zendo,  a  river,  15  m  wide,  flowing  from  the  Anko  hill  eastwards  to  the  Maze  river,  an  affluent 

of  the  Omo.     Collecting  altitude  4300  feet. 
Zuga,  a  river,  30  m  wide,  running  from  the  hill  Marta  to  Lake  Stephanie.     Collecting  altitude 

4200  feet. 

APPENDIX  3 

Barbus  susanae,  a  replacement  name  for  Barbus  gregorii  Norman. 

Barbus  gregorii  Norman  (1923)  from  China  is  preoccupied  by  Barbus  gregorii 
Boulenger  (1902)  from  Africa.  Fowler  (1958)  noticed  this  and  proposed  Barbus 
yunnanensis  to  replace  Barbus  gregorii  Norman.  Fowler  had  unfortunately  over- 
looked the  fact  that  Barbus  yunnanensis  had  already  been  used  by  Regan  (1904) 
for  a  fish  from  Yunnan,  China,  the  same  area  as  that  from  which  Barbus  gregorii 
Norman  comes. 

I  have  examined  the  type  of  Barbus  yunnanensis  Regan  B.M.  (N.H.)  Reg.  No. 
1904.1.26  :  78  and  the  syntypes  of  Barbus  gregorii  Norman  B.M.  (N.H.)  Reg.  Nos. 
1923.2.21  :  29-36,  and  they  are  very  distinct. 

I  therefore  propose  Barbus  susanae  to  replace  Barbus  gregorii  Norman. 

Derivation  of  name  :   named  after  my  wife. 

BOULENGER,  G.  A.,  1902 

Description  of  new  fishes  from  the  collection  made  by  Mr  E.   Degen  in   Abyssinia. 

Ann.  Mag.  nat.  Hist.  (7)  10  :  422. 
REGAN,  C.  T.,  1904 

A  collection  of  fishes  made  by  Mr  J.   Graham  at  Yunan  Fu.     Ann.  Mag.  nat.  Hist. 

(7)  13  :  i9i. 
NORMAN,  J.  R.,  1923 

Three  new  fishes  from  Yunan  collected  by  Professor  J.  W.  Gregory,  F.R.S.     Ann.  Mag. 

nat.  Hist.  (9)  2  :  562. 
FOWLER,  H.  W.,  1958 

Some  new  taxonomic  names  for  fish-like  vertebrates.     Not.  nat.  (310).     August  1958  :  12. 

APPENDIX  4 

A  complete  list  of  the  registered  material  examined. 

In  some  cases  more  specimens  were  examined  than  were  used  in  the  description 
of  the  species,  consequently  a  complete  list  was  deemed  advisable. 

The  Ethiopian  Barbus  spp.  from  the  Stazione  Idrobiologia,  Rome,  do  not  have 
any  register  numbers. 

The  species  are  arranged  alphabetically  and  the  following  code  applies  : 

B.M.  (N.H.)  British  Museum  (Natural  History) 

C.F.M.  Chicago  Field  Museum 

M.A.C.T.  Musee  d'Afrique  Centrale,  Tervuren 

M.H.N.P.  Museum  National  d'Histoire  Naturelle,  Paris 

M.S.N.G.  Museo  Civico  di  Storia  Naturale  G.  Doria,  Genova 

P.A.S.  Philadelphia  Academy  of  Natural  Sciences,  U.S.A. 

S.M.F.  Senckenberg  Museum,  Frankfurt 

S.M.N.H.  Stockholm  Museum  of  Natural  History 

Z.M.B.  Zoologisches  Museum  an  der  Humboldt  -  Universitat  zu  Berlin 

Z.M.H.  Zoologisches  Staatsinstitut  und  Zoologisches  Museum,  Hamburg. 


I44  K-    E-    BANISTER 

The  first  paragraph  contains  the  register  numbers  of  the  type  specimens  of  the 
species  and  of  the  species  now  in  the  synonymy.  The  following  paragraph  contains 
entirely  non-typical  material. 

Barbus  acuticeps  M.A.C.T.  130313  (holotype)  ;    130310-312  ;    130314  (paratypes). 

M.A.C.T.  91755-6,  92214. 

Barbus  alluaudi  M.H.N.P.  09-586,  09-587  (syntypes). 

Barbus    altianalis    B.M.  (N.H.)    1906.9.6  :  13    (lectotype    of    Barbus    altianalis    altianalis) 
1906.9.7  :  41   (holotype  of  Barbus  altianalis  eduardianus)  ;    1904.5.19  :  13  (holotype  of  Barbus 
altianalis  radcliffii)  ;    1906.5.30  :  117-121  ;    1906.9.6  :  14-15  ;    1906.9.7  :  41  ;    1906.9.7  :  42-43 
1911.3.3:5;     1929.1.24:84;    1929.1.24:105-108;     1929.1.24:191-192;    M.H.N.P.   30-118 
35-153  ;    Z.M.H.  19052. 

B.M.  (N.H.)  1906.5.30:107-115;  1912.10.15:15-19;  1912.10.31:1;  1925.8.7:1-2 
1928.1.25  :  12  ;  1928.5.24  :  18-25  '•  1932.6.13  :  280-289  ;  I932-6.i3  :  312-320  ;  1932.6.13  :  332 
1938.12.6:19;  1957.10.1:1-9;  1962.7.26:1-14;  1965.11.7:7-10;  1966.6.3:192;  1966.9.1 
i;  1966.9.1:2-4;  1967.5.17:52-60;  1971.1.5:121-133;  1971.2.19:28-57;  1971.8.31 
7-12  ;  1971.8.31  :  28-39  ;  M.H.N.P.  35-143,  144  ;  M.A.C.T.  66182  ;  66183  ;  92608  ;  91118- 
91122;  129096;  Z.M.H.  19052. 

Barbus  bynni  B.M.  (N.H.)  1907.12.2  :  1230  (neotype)  ;  1932.6.13  :  300-303  ;  M.S.N.G. 
J7333;  M.H.N.P.  05-275. 

B.M.  (N.H.)  1861.9.9  :  39-43  ;  1861.9.9  :  64  ;  1862.6.17  :  117-122  ;  1862.6.17  :  130-131  ; 
1905.10.26:11;  1907.12.2:1181-1229;  1907.12.2:1231-1251;  1907.12.2:3721-3728; 
1908.1.20:111-116;  1908.11.7:1;  1929.1.24:109-112;  1937.4.20:7-9;  1968.7.24:37; 
1970.12.15  :  i  ;  M.S.N.G.  17337  ;  P.A.S.  16710. 

Barbus  caudovittatus  M.A.C.T.  1168  (holotype)  ;  B.M.  (N.H.)  1901.12.26  :  26  (paratype)  ; 
1919.7.24  :  7  ;  1919.7.24  :  8-9. 

B.M.  (N.H.)  1920.5.26  :  75-76  ;  1936.6.15  :  643-644  ;  M.A.C.T.  6785  ;  6786  ;  6992  ;  14551  ; 
154!°;  15584;  39456;  43823;  44446-44461;  44482;  44483;  44551-44563;  47341; 
48504;  48505;  50061-62;  56416;  56417;  61304-63011;  74754;  774°7;  78927;  78928; 
81618;  81619;  81620;  81622-31;  81632-35;  81637-81656;  81661-81984;  81988;  91117; 
92561  ;  92562  ;  92563-78  ;  92608  ;  94318  ;  96108-114  ;  99654  ;  102024  ;  121781-826  ; 
124937  ;  125774-780  ;  129095  ;  130067  ;  130068  ;  131355  ;  134956 ;  134957  ',  ^8957  ', 
148829  ;  148830  ;  148839  ;  153485  ;  160152-163  ;  160165-67  ;  164571 ;  166954. 

Barbus  ethiopicus  B.M.  (N.H.)  1971.7.12  :  1-3. 

Barbus  gananensis  M.S.N.G.  17525  (holotype,  not  seen  by  me)  ;    M.S.N.G.  17331  ;    17339  ; 

I7341 ;  17342. 

Barbus  intermedius  S.M.F.  6778  (holotype  of  Barbus  intermedius  intermedius)  ;    B.M.  (N.H.) 
No.  18  in  1932.6.13  :  191-200  (holotype  of  Barbus  intermedius  australis)  ;    S.M.F.  2586  ;    2619  ; 
6779  ;     6786  ;     M.H.N.P.    05-252  ;     05-257  ;     B.M.  (N.H.)    1893.12.2  :  40-45  ;     1902.12.13 
211-212;     1902.12.13:225-228;     1902.12.13:261-270;     1902.12.13:274-275;     1902.12.13 
294;     1902.12.13:295-298;     1902.12.13:303-304;    1902.12.13:305-306;     1902.12.13:309 
1902.12.13:328-331;     1903.11.16:1-7;     1908.1.20:100;     1908.1.20:103-106;     1908.1.20 
107-109;       1908.1.20:110;       1908.1.20:131-133;       1908.1.20:170;     1908.1.20:181-183 
1916.1.14  :  7  ;    1937.4.20  :  66. 

B.M.  (N.H.)  1893.12.2  :  46-47  ;  1901.6.24  :  83-85  ;  1902.12.13  :  229  ;  1902.12.13  :  231-250 
1902.12.13:277-283;  1902.12.13:284-290;  1902.12.13:308;  1902.12.13:311;  1902.12.13 
312-315  ;  1902.12.13  :  317-319  ;  1902.12.13  :  320-326  ;  1902.12.13  :  332-337  ;  1902.12.13 
339;  1902.12.13  :  357 ;  1908.1.20:86;  1908.1.20:91-95;  1908.1.20:97-99;  1908.1.20 
101-102  ;  1908.1.20  :  117-130  ;  1908.1.20  :  134-155  ;  1908.1.20  :  157-168  ;  1908.1.20 
171-180;  1912.3.22:50-60;  1912.11.11:9-10;  1932.6.13:191-200  (less  the  holotype  o: 
Barbus  intermedius  australis);  1932.11.5:246-60;  1937.4.20:29-37;  1937.4.20:39-60 


A   REVISION   OF  THE   LARGE   AFRICAN   BARBUS  145 

1937.4.20:65;      1959.12.15:82-86;      1968.7.24:3-7;      1968.7.24:9-17;      1968.7.24:20-48; 
1971.8.31:15-16;    P.A.S.  14541  ;    14542. 

Barbus  longifilis  M.H.N.P.  35-150  (lectotype)  ;    35-145  -  149  ;    35~I5I  >    35-^5^- 
M.H.N.P.  35-67  ;    M.A.C.T.  42934. 

Barbus  macrolepis  Z.M.H.  £[331  (lectotype). 

B.M.  (N.H.)  1909.2.25  :  8  ;    1922.4.18  :  13  ;    1971.6.22  :  131-134.  1972-11.28  :  9-12. 

Barbus  mariae  holotype  not  seen. 

B.M.  (N.H.)  1936.12.22  :  35-39  ;    iQ37-6-4  :  l6- 

Barbus  microbarbis  M.A.C.T.  41847  (holotype). 

Barbus  microterolepis  B.M.  (N.H.)  1902.12.13  :  220  (holotype). 

Barbus  mirabilis  Z.M.B.  19059  (holotype). 

Barbus  oxyrhynchus  Z.M.H.  H339  (lectotype)  ;  B.M.  (N.H.)  1893.12.2  :  24-29  ;  1893.12.2 
32-34  ;  1893.12.2  :  37-39  ;  1902.5.26  :  25-28  ;  1902.5.26  :  35-38  ;  1906.8.25  :  7-9  ;  M.H.N.P 
26-285  ;  S.M.N.H.  9238  ;  F.M.C.  6108  ;  6109  ;  Z.M.H.  340. 

B.M.  (N.H.)  1893.12.2  :  30  ;  1893.12.2  :  35  ;  1902.5.26  :  29-34  •  I9°4-I-3°  :  5-18  ;  1904.10.10 
2-4;  1904.12.23:48-49;  1906.8.25:6;  1906.8.25:10-16;  1908.9.17:8-12;  1909.11.15 
43-47;  1914.9.21:4;  1915.12.2:2-20;  1936.12.22:2;  1936.12.22:30-34;  1937.12.11 
1-4;  1937.6.4:15;  1959.12.15:81;  1965.11.1:7-12;  1966.6.3:252;  1969.3.24:10;  1969.3.24 
44-50  ;  1971.5.10  :  55  ;  1971.8.12  :  2-6  ;  F.M.C.  6110  ;  M.A.C.T.  47341  ;  S.M.N.H.  8061 

Barbus  pagenstecheri  Z.M.H.  H.342  (lectotype),  H.34I  (paralectotype) . 
Barbus  paucisquamatus  M.H.N.P.  35-76  (lectotype). 

M.H.N.P.  35-77;  35-78;  35-I53:  35-r54  '.  35-IlS  (3  specimens);  M.A.C.T.  42932; 
130145  ;  130146. 

Barbus  platyrhinus  B.M.  (N.H.)  1906.9.6  :  12  (holotype). 
M.A.C.T.  89789-92  ;    130658-60. 

Barbus  ruasae  Z.M.B.  19053  (lectotype,  one  of  two  specimens). 
M.A.C.T.  91755 ;   91756  ;   92214. 

Barbus  somereni  B.M.  (N.H.)  1911.7.26  :  i  (holotype)  ;    M.A.C.T.  46963-65. 

B.M.  (N.H.)  1969.3.3:13-14;  1971.1.5:96-99;  1971.1.5:100-117;  1971.1.5:120; 
1971.1.5:145;  1971.2.19:26-27;  1971.8.31:13-14;  1971.10.18:1-5;  M.A.C.T.  46952-62  ; 
46966-47339;  47342;  55788-792;  56456-478;  56480-91;  71781-82;  73I57~73i62  ; 
87692-695;  87696-698;  94221-232;  92579-89;  92591-92607;  93356-377;  129108. 

Barbus  stappersii  M.A.C.T.  14250  (holotype)  ;    14765  ;    14233  ;    14172. 
M.A.C.T.  81945  ;    81985  ;    122295  '•    !38958-959. 

Barbus  trachypterus  M.A.C.T.  81621  (holotype). 
M.A.C.T.  129097-100  ;    165254-256. 

Barbus  tropidolepis  B.M.  (N.H.)   1906.9.6:  19-21    (the  smallest  specimen  is  the  lectotype). 

B.M.  (N.H.)     1906.9.8  :  50-52  ;       1920.5.25  :  38-46  ;       1936.6.15  :  568-629  ;       1955.12.20  : 

720-731;    1955.12.20:733-815;    1955.12.20:873;    1955.12.20:888;    1955.12.20:1169-1171. 

INDEX 

The  generic  name  Barbus  has  been  omitted  from  this  list.  Where  the  genus  is  not  Barbus  it  is 
Varicorhinus  and  is  symbolized  by  V. 

acuticeps  Matthes  1959,  5 

affinis  Ruppell  1837  =  B.  intermedius  intermedius,  47 

affinis  brevibarbus  Bini  1940  =  B.  intermedius  intermedius,  47 

affinis  nedgia  Bini  1940  =  B.  intermedius  intermedius,  47 

ahlselli  Lonnberg  1911  =  B.  oxyrhynchus,  94 


146  INDEX 

alluaudi  Pellegrin  1909,  9 

altianalis  Blgr.  1900,  9 

altianalis  altianalis  Worthington  1932,  9 

altianalis  eduardianus  Worth.  1932,  9 

altianalis  radcliffii  Worth.  1932,  9 

altianalis  var.  labiosa  Pell.  1932  =  B.  altianalis  altianalis,  g 

altianalis  var.  lobogenysoides  Pell.  1935  =  B.  paucisquamatus,  102 

altianalis  var.  longifilis  Pell.  1935  =  B.  longifilis,  76 

altianalis  var.  paucisquamata  Pell.  1935  =  B.  paucisquamatus,  102 

altianalis  urundensis  David  1937  =  &•  somereni,  in 

alticola  Blgr.  1906  =  B.  intermedius  intermedius,  47 

a^Ai  Hubbs  1918  =  B.  oxyrhynchus,  94 

babaulti  Pell.  1926  =  B.  oxyrhynchus,  94 

babaulti  Pell.  1935  =  B.  lapsus,  94 

F.  babaulti  :   Bertin  and  Esteve  1947  =  B.  oxyrhynchus,  94 

bayoni  Blgr.  1911  =  B.  altianalis  altianalis,  9 

Capoeta  bingeri  Pell.  1905  =  B.  intermedius  intermedius,  47 

bingeri  :   Blgr.  1911  =  .B.  intermedius  intermedius,  47 

F.  bingeri  :   Bertin  and  Esteve  1947  =  B.  intermedius  intermedius,  47 

bottegi  Blgr.  1906  =  B.  intermedius  intermedius,  47 

F.  bredoi  Poll  1948  =  B.  trachypterus,  119 

brevibarbus  Blgr.  1902  =  B.  intermedius  intermedius,  47 

brunellii  Bini  1940  =  B.  intermedius  intermedius,  47 

brunellii  acutirostris  Bini  1940  =  B.  intermedius  intermedius,  47 

bynni  (Forssk.)  1775,  27 

bynni  rudolfianus  Worthington  1932  =  B.  bynni,  27 

caudovittatus  Blgr.  1902,  36 

F.  chapini  Nicholls  and  LaMonte  1950  =  B.  tropidolepis,  123 

chilotes  sakaniae  Poll  1938  =  B.  caudovittatus,  36 

curtus  Blgr.  1915  =  B.  stappersii,  116 

dainellii  Bini  1940  =  B.  intermedius  intermedius,  47 

dainellii  macrocephalus  Bini  1940  =  B.  intermedius  intermedius,  47 

degeni  Blgr.  1902  =  B.  intermedius  intermedius,  47 

degeni  leptorhinus  Bini  1940  =  B.  intermedius  intermedius,  47 

donyensis  Holly  1929  =  B.  oxyrhynchus,  94 

duchesnii  Blgr.  1902  =  B.  intermedius  intermedius,  47 

duchesnii  ibridus  Bini  1940  =  B.  intermedius  intermedius,  47 

duchesnii  maximus  Bini  1940  =  B.  intermedius  intermedius,  47 

eduardianus  Blgr.  1901  =  B.  altianalis  eduardianus,  9 

elongatus  Riipp.  1837  =  B.  intermedius  intermedius,  47 

erlangeri  Blgr.  1903  =  B.  intermedius  intermedius,  47 

ethiopicus  Zolezzi  1940,  41 

euchilus  Blgr.  1919  =  B.  caudovittatus,  36 

eumystus  Blgr.  1906  =  B.  intermedius  intermedius,  47 

fergussonii  Blgr.  1901  =  B.  altianalis  eduardianus,  9 

gananensis  Vincig.  1895,  44 

gorguarii  Riipp.  1837  =  B.  intermedius  intermedius,  47 

gorguarii  macroptalmus  Bini  1940  =  B.  intermedius  intermedius,  47 


INDEX  147 


gregorii  Blgr.  1902  =  B.  intermedius  australis,  75 
gudaricus  Blgr.  1906  =  B.  intermedius  intermedius,  47 

harringtonii  Blgr.  1902  =  B.  intermedius  intermedius,  47 
hindii  Blgr.  1902  =  B.  oxyrhynchus,  94 
hollyi  Lohberger  1929  =  B.  altianalis  radcliffii,  g 
hursensis  Blgr.  1902  =  B.  intermedius  intermedius,  47 

ilgi  Pell.  1905  =  B.  intermedius  intermedius,  47 

intermedius  Rtipp.  1837,  47 

intermedius  australis  s.  sp.  nov.,  75 

intermedius  gorgorensis  Bini  1940  =  B.  intermedius  intermedius,  47 

intermedius  intermedius  s.  sp.  nov.,  74 

intermedius  microstoma  Bini  1940  =  B.  intermedius  intermedius,  47 

jarsinus  Blgr.  1902  =  B.  intermedius  intermedius,  47 

kassamensis  Blgr.  1902  =  B.  intermedius  intermedius,  47 
kiogae  Worth.  1929  =  B.  altianalis  radcliffii,  9 
kivuensis  Pappenheim  1914  =  B.  altianalis  altianalis,  9 
krapfi  Blgr.  1911  =  B.  oxyrhynchus,  94 

labiatus  Blgr.  1902  =  B.  oxyrhynchus,  94 

leptosoma  Blgr.  1902  =  B.  intermedius  intermedius,  47 

lestradei  David  1936  =  B.  caudovittatus,  36 

lobogenys  Blgr.  1906  =  B.  altianalis  radcliffii,  9 

longifilis  :   Pellegrin  1935,  ?6 

longirostris  Worth.  1929  =  B.  altianalis  radcliffii,  9 

macmillani  Blgr.  1906  =  B.  intermedius  intermedius,  47 

macrolepis  Pfeffer  1889,  79 

macronema  Blgr.  1902  =  B.  intermedius  intermedius,  47 

macronema  var.  parenzani  Zolezzi  1940  =  B.  intermedius  intermedius,  47 

margaritae  Blgr.  1906  =  B.  intermedius  intermedius,  47 

mariae  Holly  1929,  83 

mathoiae  Blgr.  1911  =  B.  oxyrhynchus,  94 

matris  Holly  1928,  83 

meneliki  Pell.  1905  =  B.  bynni,  27 

mento  Blgr.  1902  =  B.  intermedius  intermedius,  47 

microbarbis  David  and  Poll  1937,  87 

microterolepis  Blgr.  1902,  89 

miochilus  Blgr.  1919  =  B.  caudovittatus,  36 

mirabilis  Papp.  1914,  91 

moeruensis  Pell.  1922  =  B.  stappersii,  116 

nairobi  Holly  1928  =  B.  oxyrhynchus,  94 

nedgia  Riipp.  1837  =  B.  intermedius  intermedius,  47 

neuvillii  Pell.  1905  =  B.  intermedius  intermedius,  47 

obesus  Worth.  1929  =  B.  altianalis  radcliffii,  9 
oreas  Blgr.  1902  =  B.  intermedius  intermedius,  47 
oxycephalus  Blgr.  1915  =  B.  stappersii,  116 
oxyrhynchus  Pfeffer  1889,  94 


148  INDEX 

pagenstecheri  Fischer  1884,  101 

paucisquamatus  :   Pellegrin  1933,  103 

perplexicans  Blgr.  1902  =  B.  oxyrhynchus,  94 

pietschmanni  Lohberger  1929  =  B.  altianalis  vadcliffii,  9 

plagiostomus  Blgr.  1902  =  B.  intermedius  intermedius,  47 

platyrhinus  Blgr.  1900,  106 

platystomus  Blgr.  1902  =  B.  intermedius  intermedius,  47 

platystomus  daga  Bini  1940  =  B.  intermedius  intermedius,  47 

platystomus  dekkensis  Bini  1940  =  B.  intermedius  intermedius,  47 

platystomus  platystomus  Bini  1940  =  B.  intermedius  intermedius,  47 

platystomus  prognathus  Bini  1940  =  B.  intermedius  intermedius,  47 

platystomus  var.  vatovae  Zolezzi  1940  =  B.  intermedius  intermedius,  47 

pojeri  Poll  1944  =  B.  caudovittatus,  36 

procatopus  Blgr.  1914  =  B.  intermedius  australis,  75 

radcliffii  Blgr.  1903  =  B.  altianalis  radcliffii,  9 

rhinoceros  Copley  1938  =  B.  mariae,  83 

ruasae  Pappenheim  and  Blgr.  1914,  109 

ruepelli  Blgr.  1902  =  B.  intermedius  intermedius,  47 

ruspolii  Vincig.  1896  =  B.  bynni,  27 

somereni  Blgr.  1911,  in 

stappersii  Blgr.  1915,  116 

V.  stappersii  Blgr.  1917  =  B.  caudovittatus,  36 

surkis  Riipp.  1837  =  B.  intermedius  intermedius,  47 


tanensis  Giinther  1894  =  B.  oxyrhynchus,  94 
trachypterus  Blgr.  1915,  119 
tropidolepis  Blgr.  1900,  123 

urundensis  :   Poll  1946  =  B.  somereni,  in 

volpinii  Parenzan  1940  =  B.  intermedius  intermedius,  47 

zaphiri  Blgr.  1906  =  B.  intermedius  intermedius,  47 
zuaicus  Blgr.  1906  =  B.  intermedius  intermedius,  47 


DR  K.  E.  BANISTER 

Department  of  Zoology 

BRITISH  MUSEUM  (NATURAL  HISTORY) 

CROMWELL  ROAD 

LONDON  SW7  560 


A  LIST  OF  SUPPLEMENTS 
TO  THE  ZOOLOGICAL  SERIES 

OF  THE  BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 


1.  KAY,  E.  ALISON.     Marine  Molluscs  in  the  Cuming  Collection  British  Museum 
(Natural  History)  described  by  William  Harper  Pease.     Pp.  96  ;    14  Plates. 
1965.     (Out  of  Print.)     £3.75.* 

2.  WHITEHEAD,  P.  J.  P.    The  Clupeoid  Fishes  described  by  Lacepede,  Cuvier  and 
Valenciennes.     Pp.  180  ;  u  Plates,  15  Text-figures.     1967.    £4. 

3.  TAYLOR,  J.  D.,   KENNEDY,  W.  J.   &  HALL,  A.    The  Shell  Structure  and 
Mineralogy  of  the  Bivalvia.     Introduction.     Nuculacea-Trigonacea.     Pp.  125  ; 
29  Plates,  77  Text-figures.     1969.     £4.50. 

4.  HAYNES,  J.  R.     Cardigan  Bay  Recent  Foraminifera  (Cruises  of  the  R.V.  Antur) 
1962-1964.     Pp.  245  ;  33  Plates,  47  Text-figures.     1973.    £10.80. 


Printed  in  Great  Britain  by  John  Wright  and  Sons  Ltd.  at  The  Stonebridge  Press,  Bristol  BS4  jNU 


SOME  DIGENETIC  TREMATODES 
FISHES  FROM  THE  BAY  OF  BISCAY 
AND  NEARBY  WATERS 


R.  A.  BRAY 


BULLETIN  OF 

THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 
ZOOLOGY  Vol.  26  No.  2 

LONDON:   1973 


SOME  DIGENETIC  TREMATODES  IN  FISHES 

FROM  THE  BAY  OF  BISCAY 

AND  NEARBY  WATERS 


BY 

RODNEY  ALAN  BRAY 


X 


Pp  149-183  ;   ii  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
ZOOLOGY  Vol.  26  No.  2 

LONDON:  1973 


THE     BULLETIN      OF     THE      BRITISH     MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they 
become  ready.  Volumes  will  contain  about  three  or 
four  hundred  pages,  and  will  not  necessarily  be 
completed  within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  26,  No.  2,  of  the  Zoological  series. 
The  abbreviated  titles  of  periodicals  cited  follow  those 
of  the  World  List  of  Scientific  Periodicals. 


World  List  abbreviation  : 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


Trustees  of  the  British  Museum  (Natural  History),  1973 


TRUSTEES  OF 
THE   BRITISH   MUSEUM    (NATURAL   HISTORY) 

Issued  15  November,  1973  Price  £1.60 


SOME  DIGENETIC  TREMATODES  IN  FISHES 

FROM  THE  BAY  OF  BISCAY 

AND  NEARBY  WATERS 

By  RODNEY  A.  BRAY 

CONTENTS 


SYNOPSIS           ...........  151 

INTRODUCTION            ..........  151 

HOST  -  PARASITE  LIST         .          .                   .          .          .          .          .          .  154 

SYSTEMATIC  SECTION           .........  155 

Family  BUCEPHALIDAE          ........  155 

Family  OPECOELIDAE  .........  159 

Family  LEPOCREADIIDAE      ........  162 

Family  AZYGIIDAE       .........  167 

Family  ZOOGONIDAE    .........  167 

Family  FELLODISTOMIDAE    .          .          .          .          .          .          .          .173 

Family  HEMIURIDAE   ........          .178 

ACKNOWLEDGMENTS  ..........  181 

REFERENCES  181 


SYNOPSIS 

Sixteen  species  of  digenetic  trematodes  are  recorded  from  seventeen  species  of  fishes  caught  in 
the  Bay  of  Biscay  and  in  waters  off  the  north-west  coast  of  Spain  and  the  Atlantic  coast  of 
Morocco. 

The  following  two  species  are  described  as  new  :  Bathycreadium  biscayense  and  Steringophorus 
blackeri.  Seven  of  the  remaining  species  are  described,  these  are  Dolichoenterum  longissimum 
Ozaki,  Prosorhynchus  crucibulum  (Rud.),  Lepidapedon  rachion  (Cobbold),  L.  elongatum  (Lebour), 
Steganoderma  abyssorum  (Odhner),  Neosteganoderma  glandulosum  Byrd  and  (?)  Steringotrema 
diver  gens  (Rud.).  The  following  new  combinations  are  made  :  Bathycreadium  [Nicolla]  elonga- 
tum (Maillard)  and  Neosteganoderma  [Proctophantastes]  polymixiae  (Yamaguti). 

INTRODUCTION 

THE  author  was  given  the  opportunity  in  1971  to  obtain  parasitic  worms  from  marine 
fishes,  whilst  accompanying  a  cruise  of  the  Ministry  of  Agriculture,  Fisheries  and 
Food  Research  Vessel  'Cirolana',  the  primary  purpose  of  which  was  to  collect  blood 
from  hake  for  electrophoretic  and  chromosome  studies.  The  collecting  was  carried 
out  between  the  ist  and  the  i4th  of  January  and  was  conducted  primarily  in  the 
Bay  of  Biscay,  but  also  in  other  areas,  including  the  waters  off  the  western  coasts  of 
Spain  and  Morocco.  The  locations  of  the  stations  mentioned  in  this  report  are 
shown  on  Fig.  i  and  further  details  are  given  in  Table  I. 

The  fish  were  caught  by  stern  trawl  and  the  alimentary  tract  removed  from  them. 
The  stomach  and  intestine  were  separated  before  being  opened  up  with  scissors,  and 
then  the  contents  were  shaken  into  a  tube  of  sea-water.  These  tubes  were  kept  for 


152 


R.   A.    BRAY 


N.I.O. 
6429 


FIG.  i.     Map  showing  positions  of  stations  mentioned  in  this  report. 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  153 


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R.   A.    BRAY 


periods  of  up  to  six  hours  in  a  refrigerator  at  just  above  freezing  point,  before  being 
topped  up  with  a  fixative,  either  'Dowicil'  or  10  per  cent  formalin.  A  degree  of 
contraction  resulted  from  the  use  of  these  fixatives. 

The  material  from  the  'Cirolana'  cruise  was  supplemented  by  two  trematodes 
collected  from  preserved  fish  in  the  collection  of  the  National  Institute  of  Oceano- 
graphy (now  the  Institute  of  Oceanographic  Sciences)  at  Wormley,  Surrey. 

The  parasites  of  seventeen  species  of  fish  are  mentioned  in  this  report .  The  maj  ority 
of  the  sixteen  species  of  digenetic  trematodes  studied  are  already  well  known  as 
widespread  species.  It  is  interesting  to  note,  however,  that  Neosteganoderma 
glandulosum,  reported  originally  from  Florida,  has  now  been  recovered  from  off  the 
north-west  coast  of  Spain  and  that  Steganoderma  abyssorum,  previously  reported 
only  from  Norway,  occurs  in  the  eastern  Bay  of  Biscay.  Further  evidence  of  the 
affinities  of  species  in  the  area  under  discussion  with  Florida  on  one  hand,  and  the 
north-eastern  North  Atlantic  on  the  other,  is  indicated  by  the  finding  of  two  new 
species  whose  closest  relatives  appear  to  be  species  from  these  two  areas. 


HOST-PARASITE   LIST 

Alosa  fallax  (Lacepede)  -  Hemiurus  appendiculatus  (Rud.). 
Beryx  decadactylus  Cuvier  -  Neosteganoderma  glandulosum  Byrd. 
Capros  aper  (L.)  -  (?)  Steringotrema  diver  gens  (Rud.). 
Coelorinchus  caelorhinchus  (Risso)  -  Bathycreadium  biscayense  sp.  nov. 

Steganoderma  abyssorum  (Odhner). 
Conger  conger  (L.)  -  Dolichoenterum  longissimum  Ozaki. 
Prosorhynchus  crucibulum  (Rud.). 
Lecithochirium  gravidum  Looss. 

Dalatias  licha  (Bonnaterre)  -  Otodistomum  veliporum  (Creplin) . 
Gaidropsarus  vulgaris  (Cloquet)  -  Helicometra  fasciata  (Rud.). 
Lepidion  eques  (Giinther)  -  Bathycreadium  biscayense  sp.  nov. 

Lepidapedon  elongatum  (Lebour). 
Steganoderma  abyssorum  (Odhner). 

Lophius  piscatorius  L.  -  Bucephahpsis  gracilescens  (Rud.). 
Merluccius  merluccius  (L.)  -  Derogenes  various  (Muller). 
Micromesistius  poutassou  (Risso)  -Derogenes  various  (Muller). 
Molva  macrophthalma  (Rafinesque-Schmaltze)  -Derogenes  various  (Muller). 
Phycis  blennoides  (Brunnich)  -  Lepidapedon  rachion  (Cobbold) . 

Derogenes  various  (Muller). 

Scomber  scombrus  L.  -  Lecithocladium  excisum  (Rud.). 
Trachyrhynchus  trachyrincus  (Risso)  -  Bathycreadium  biscayense  sp.  nov. 

Lepidapedon  elongatum  (Lebour). 
Steganoderma  abyssorum  (Odhner). 
Derogenes  various  (Muller). 

Xenodermichthys  copei  (Gill)  -  Steringophorus  blackeri  sp.  nov. 
Xenodermichthys  sp.  indet.  -  Steringophorus  blackeri  sp.  nov. 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  155 

SYSTEMATIC   SECTION 

Family  BUCEPHALIDAE  Poche,  1907 

Dolichoenterum  longissimum  Ozaki,  1924 

(Fig.  2) 

Host  and  locality  :   intestine  of  Conger  conger,  Stn.  7. 

Twenty-three  specimens  were  found,  and  these  vary  from  3-5  to  6-5  mm  in  length. 
The  longer  worms  are  less  contracted  and  have  a  long  narrow  neck,  and  an  elongate 
oval  hindbody  (Fig.  2a),  but  in  the  shorter  contracted  specimens  there  is  little  indica- 
tion of  a  neck  (Fig.  2b).  Whilst,  superficially,  these  two  forms  may  appear  distinct 
an  examination  of  the  internal  morphology  of  the  worms  leaves  no  doubt  that  they 
are  specifically  identical.  The  cuticle  has,  imbedded  in  it,  numerous  narrow  pointed 
spines. 

The  anterior  sucker  is  surmounted  by  a  thick  muscular  hood,  which  in  an  extended 
condition,  bears  seven  to  eight  horn-like  conical  projections  on  the  dorsal  and  lateral 
rims  (Figs.  2c  and  d).  In  an  extended  condition  the  sucker  is  0-53-0-68  mm  in 
diameter.  When  contracted  the  sucker  becomes  almost  globular,  and  the  projections 
are  less  conspicuous,  and  apparently  less  numerous  in  some  cases,  and  point  inwards. 
The  sucker  in  this  condition  measures  0-32-0-44  mm  in  diameter.  The  aperture  of 
the  sucker  is  subterminal,  with  a  more  or  less  narrow  extension  down  the  ventral 
surface. 

The  contraction  or  expansion  of  the  worm  also  affects  the  anterior  portion  of  the 
alimentary  canal.  In  neither  whole  mounts  nor  serial  sections  was  evidence  of  an 
oral  sucker  seen.  The  specimens  lacking  a  neck  bear  a  simple  mouth  situated  closely 
behind  the  anterior  sucker.  A  short  prepharynx  runs  posteriorly  to  the  somewhat 
transversely-oval  pharynx  measuring  0-3-0-32  mm  x  0-21  mm.  From  the  pharynx 
the  intestine  runs  anteriorly  to  a  point  which  may  or  may  not  be  nearer  to  the  an- 
terior sucker  than  the  pharynx,  where  it  turns  conversely  to  reach  the  hinder  end 
of  the  body.  In  both  extended  and  contracted  specimens  the  intestine  follows  a 
transversely  undulating  course  and  reaches  nearly  to  the  posterior  end  of  the  cirrus- 
sac.  The  posterior  limit  of  the  intestine  is  not  easy  to  make  out  in  whole  mounts  as 
it  is  often  obscured  by  the  eggs  in  the  uterus. 

A  sinus  occurs  at  the  posterior  margin  of  the  worm,  normally  in  the  median  line, 
though  contraction  has,  in  some  cases,  displaced  the  opening  to  the  left  of  the 
median  line.  Into  it  open  the  ducts  of  the  male  and  female  systems  and  of  the  excre- 
tory vesicle.  The  opening  of  the  sinus  does  not  appear  to  be  surrounded  by  a 
sphincter.  The  elongate  cirrus-sac,  which,  due  to  contraction,  lies  at  an  acute  angle 
to  the  median  line,  measures  0-95  x  0-23-0-28  mm  and  possesses  a  fairly  thick  mus- 
cular wall.  It  contains  an  oval  seminal  vesicle,  0-38-0-43  mm  x  0-23-0-25  mm, 
which  occupies  the  anterior  portion  of  the  cirrus-sac.  The  pars  prostatica  passes 
out  of  the  middle  region  of  the  seminal  vesicle,  and  runs  to  near  the  posterior  ex- 
tremity of  the  cirrus-sac  where  it  unites  with  a  short  muscular  cirrus.  The  testes  are 
transversely  elongate,  well  separated  one  behind  the  other,  and  are  situated  in  the 


R.    A.    BRAY 


FIG.  2.  Dolichoenterum  longissimum  Ozaki  :  a)  extended  specimen  (ventral  view)  ; 
b)  contracted  specimen  (ventral  view)  ;  c)  anterior  sucker  (dorso-lateral  view)  ;  d) 
longitudinal  section  of  anterior  sucker. 


middle  region  of  the  hinder  half  of  the  body.  They  are  0-56-0-84  mm  x  0-22-0-41 
mm.  The  ovary  is  also  transversely  elongate  measuring  0-53-0-82  mm  x  0-23- 
0-34  mm,  and  lies  a  little  to  the  left  of  the  median  line  between  the  testes.  Directly 
behind  it  lies  a  large  diffuse  Mehlis'  gland.  Laurer's  canal  opens  dorsally  at  about 
the  level  of  the  posterior  testis.  There  appears  to  be  no  receptaculum  seminis,  but 
sperm  is  accumulated  in  the  initial  slings  of  the  uterus,  which  is  long  and  coiled 
running  to  a  position  some  way  anterior  to  the  anterior  testis  before  it  turns  and  passes 
posteriorly,  with  many  tightly-packed  transverse  slings.  It  passes  into  the  posterior 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  157 

region  of  the  body,  where  it  opens  into  the  base  of  the  genital  sinus  near  the  cirrus- 
sac,  via  a  short  muscular  metraterm.  There  are  numerous  small  eggs,  many  of 
which  are  collapsed,  but  measuring  21-27  /mi  x  12-18  /mi  when  not  collapsed. 
The  vitelline  follicles  lie  in  two  lateral  groups,  with  about  12-16  follicles  in  each 
group,  extending  from  about  the  level  of  the  anterior  testis  to  a  position  anterior  to 
the  foremost  extent  of  the  uterus.  A  few  follicles  also  occur  scattered  between  the 
testes. 

The  excretory  vesicle  opens  into  the  genital  sinus,  and  runs  forward  to  a  position 
just  posterior  to  the  posterior  testis. 

Dolichoenterum  longissimum  was  originally  described  from  Conger  myriaster  in 
Japanese  waters.  It  has  been  found  a  number  of  times  since  in  the  same  waters  in 
C.  myriaster  (Ozaki  1928  ;  Yamaguti  1938  ;  Machida  et  al.  1970)  and  in  Muraenesox 
cinereus  (Yamaguti  1934).  Prior  to  the  present  record,  it  has  also  been  found  in 
Conger  conger  in  New  Zealand  waters  (Manter  1954)  and  in  two  localities  off  the 
western  coast  of  Scotland  (Williams  1960). 

Ozaki  (1928)  and  Yamaguti  (1934)  disagree  on  the  presence  of  a  so-called  '  urogeni- 
tal '  pore,  that  is,  the  common  opening  of  the  genital  sinus  and  the  excretory  vesicle. 
Yamaguti  states  that  Ozaki's  figure  23  is  misleading,  and  suggests  that  the  'genital 
sinus'  is  no  more  than  a  slight  depression.  Ozaki's  figure,  however,  seems  to 
accurately  represent  the  condition  in  the  present  material.  It  would  seem  that  the 
depth  of  the  genital  sinus  is  influenced  somewhat  by  the  contraction  of  the  body. 

Another  feature  which  varies  with  the  condition  of  the  worm  is  the  number  and 
configuration  of  the  projections  surmounting  the  anterior  sucker.  Such  a  difference 
is,  according  to  Tendeiro  (1955),  a  major  factor  in  separating  D.  manteri  Tendeiro, 
1955  (with  only  five  projections)  from  D.  longissimum.  Tendeiro  found  a  single 
specimen  of  D.  manteri  in  C.  conger  from  the  Algarve  coast  of  Portugal,  and  considered 
it  distinct  for  a  number  of  other  reasons.  The  only  one  of  these  that  appears  to  be 
a  valid  differentiating  feature  is  the  length  of  the  intestine,  which  in  D.  manteri  is 
said  to  extend  only  a  little  behind  the  middle  region  of  the  body.  As  Tendeiro  had 
but  one  specimen  it  may  have  been  that,  as  is  the  case  in  many  of  my  specimens,  it 
was  not  possible  to  trace  the  full  extent  of  the  intestine.  If  this  were  found  to  be  so, 
then  it  seems  probable  that  D,  manteri  should  be  considered  a  synonym  of  D. 
longissimum. 

Bucephalopsis  gracilescens  (Rud.,  1819)  Nicoll,  1914 

Host  and  locality  :  intestine  of  Lophius  piscatorius,  Stn.  54. 

This  parasite  occurred  in  great  numbers.  The  shape  of  these  worms  varies 
considerably  as  indicated  by  Dawes  (1947). 

Prosorhynchus  crucibulum  (Rud.,  1819)  Odhner,  1905 

(Fig.  3) 

Monostoma  crucibulum  Rud.,  1819. 

Distoma  crucibulum  :   Dujardin,  1845. 

Gasterostomum  crucibulum  :   Gervais  and  van  Beneden,  1859. 


158 


R.   A.    BRAY 


FIG.  3.     Prosorhynchus  crucibulum  (Rud.)  :    ventral  view. 


Host  and  locality  :  intestine  of  Conger  conger,  Stn.  7. 

A  single  specimen  was  present,  and  it  was  examined  as  a  whole  mount  and  then 
in  serial  sections.  It  is  a  small  worm,  truncated  anteriorly  and  pointed  posteriorly, 
1-5  mm  in  length  and  0-87  mm  in  maximum  width.  The  cuticle  bears  numerous 
scale-like  spines.  At  the  anterior  end  there  is  a  muscular,  funnel-shaped  rhynchus, 
0-48  mm  across  at  its  widest  point  and  extending  posteriorly  to  very  near  the  intes- 
tine, that  is,  about  0-55  mm  from  the  anterior  end  of  the  worm.  A  slit-like  transverse 
mouth,  52  /im  across,  is  situated  at  about  the  anterior  third  of  the  body  length,  and 
near  the  base  of  the  rhynchus.  It  leads  immediately  to  a  nearly  globular  pharynx, 
measuring  0-108  x  0-115  mm,  which  in  turn  leads  into  a  muscular  oesophagus 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  159 

0-07  mm  long,  thence  into  the  saccular  intestine,  which  lies  dorsally  to  the  pharynx 
and  is  about  0-2  mm  x  0-16  mm. 

The  excretory  pore  is  terminal  posteriorly,  and  the  excretory  vesicle  reaches  to 
about  the  level  of  the  anterior  end  of  the  cirrus-sac. 

The  genital  sinus  opens  near  to  the  posterior  extremity  of  the  body  on  the  ventral 
side  of  the  worm.  A  muscular  cirrus-sac,  0-47  mm  long  by  about  0-22  mm  wide, 
reaches  anteriorly  to  a  position  just  in  front  of  the  anterior  margin  of  the  posterior 
testis.  It  contains  a  sigmoid  seminal  vesicle,  which  extends  from  the  foremost 
extremity  of  the  cirrus-sac  to  a  position  near  to  the  posterior  extremity,  where  it 
forms  a  muscular  bulb  before  passing  into  an  elongate  pars  prostatica.  This  latter 
organ  runs  nearly  to  the  anterior  of  the  cirrus-sac  before  passing  posteriorly  to  where 
it  unites  with  a  relatively  short  muscular  cirrus,  which  opens  into  the  sinus  at  the  base 
of  the  genital  tongue.  There  is  no  indication  of  an  external  seminal  vesicle.  The 
testes  lie  one  just  posterior  to  the  pharynx  with  the  other  just  behind  it.  They  are 
oval  with  smooth  margins  and  measure  0-27  x  0-19  mm  (anterior  testis)  and  0-21  x 
0-2  mm  (posterior  testis).  The  ovary,  of  similar  size  and  shape  to  the  testes  (0-2  mm 
dia.),  lies  just  antero-dorsal  to  the  anterior  testis.  The  '  shell  '-gland  lies  immediately 
posterior  to  the  ovary,  and  Laurer's  canal  opens  dorsally  just  posterior  to  the  ovary. 
The  vitellaria  consist  of  about  25  irregular  follicles  lying  more  or  less  in  an  arc  just 
behind  the  rhynchus  and  anterior  to  the  pharynx  and  uterus.  This  latter  organ  is 
extensive  in  the  region  posterior  to  the  pharynx,  obscuring  parts  of  the  reproductive 
system  in  the  whole  mount.  A  short,  muscular  metraterm  enters  the  genital  sinus 
dorsally  to  the  cirrus-sac.  The  eggs,  which  are  very  numerous  and  mostly  collapsed, 
measure  about  24-26  p,m  x  16-18  p,m. 

Prosorhynchus  crucibulum  is  a  common  parasite  of  the  conger  and  is  morphologi- 
cally very  similar  to  another  parasite  of  this  fish,  P.  aculeatus  Odhner,  1905,  with 
which  it  has  been  synonymized  by  Dawes  (1947).  Brinkmann  (1957)  has  given  a 
number  of  reasons  why  he  considers  these  species  to  be  distinct,  and,  having  examined 
a  number  of  specimens  assigned  to  both  species  in  the  collections  of  the  British  Museum 
(Natural  History),  I  agree  that  the  features  selected  by  Brinkmann  appear  to  be  of 
value  in  distinguishing  these  species.  The  most  noticeable  difference  is  the  size  and 
shape  of  the  rhynchus,  which  in  P.  aculeatus  is  small  and  rounded,  and  in  crucibulum 
is  larger  and  triangular  or  funnel-shaped.  This  character  appears  to  be  fairly 
constant,  and  according  to  Matthews  (1973)  the  typical  shape  of  the  rhynchus  in 
P.  crucibulum  is  developed  in  the  metacercaria  during  its  first  month  in  the  second 
intermediate  host.  As  can  be  seen,  the  present  specimen  fits  P.  crucibulum  in  this, 
as  well  as  the  other  less  prominent  characteristics. 


Family  OPECOELIDAE  Ozaki,  1925 
H elicometra  fasciata  (Rud.,  1819)  Odhner,  1902 

Host  and  locality  :  intestine  of  Gaidropsarus  vulgaris,  Stn.  22. 
One  specimen  was  present. 


160  R.   A.    BRAY 

Bathycreadium  biscayense  sp.  nov. 
(Fig-  4) 

Hosts  and  localities  :  caeca  and  intestine  of  Trachyrhynchus  trachyrincus  (type-host), 
Stn.  7,  Stn.  24  ;  caeca  and  intestine  of  Coelorinchus  caelorhinchus,  Stn.  i  ;  intestine 
of  Lepidion  eques,  Stri.  i. 

This  species  occurred  commonly  in  T.  trachyrincus,  in  smaller  numbers  in  C. 
caelorhinchus  and  only  one  specimen  was  found  in  L.  eques. 

The  worm  is  elongate  oval  with  lateral  margins  roughly  parallel  for  much  of  its 
length,  which  varies  between  0-4  mm  and  1-7  mm.  The  maximum  width,  which 
normally  occurs  at  the  level  of  the  ventral  sucker,  is  0-19-0-42  mm.  The  tegument 
is  not  armed,  but  bears  annular  wrinkles,  probably  due  to  contraction.  Circular  and 
subterminal,  the  oral  sucker  measures  0-065-0-14  mm  in  diameter  and  leads,  without 
an  apparent  prepharynx,  into  a  globular  pharynx  0-032-0-066  mm  in  diameter. 
The  oesophagus,  up  to  0-13  mm  in  length  and  lined  with  a  cuticle,  bifurcates  at  about 
the  anterior  margin  of  the  ventral  sucker.  The  intestinal  caeca  reach  to  near  the 
posterior  end  of  the  worm,  where  they  unite  to  form  a  cyclocoel.  The  ventral  sucker 
is  situated  close  behind  the  oral  sucker  in  the  anterior  quarter  or  third  of  the  body. 
It  is  often  situated  on  a  slight  protuberance,  and  is  transversely  elongate,  its  trans- 
verse diameter  being  0-19-0-29  mm,  thus  giving  an  oral :  ventral  sucker  ratio  of 
i  :  2-2-3-5. 

The  excretory  pore  is  terminal,  and  the  cylindrical  vesicle  reaches  to  the  ovary. 

The  genital  pore  occurs  just  to  the  left  of  the  median  line,  closely  anterior  to  the 
ventral  sucker.  Into  it  opens  a  claviform  cirrus-sac  measuring  0-25-0-4  mm  x  0-05- 
0-085  mm,  which  normally  reaches  just  posteriorly  to  the  ventral  sucker,  and  its 
posterior  extremity  lies  amongst  the  uterine  coils.  It  contains  a  convoluted  internal 
seminal  vesicle,  a  relatively  short,  oval  pars  prostatica  and  a  long  muscular  cirrus. 
The  testes  are  situated  in  tandem,  close  together,  in  the  middle  third  of  the  body. 
They  are  usually  wider  than  long  with  entire  or  slightly  indented  margins  and  measure 
in  mature  specimens  0-08-0-12  mm  x  0-12-0-15  mm  (anterior  testis)  and  0-09- 
0-12  mm  x  0-12-0-15  mm  (posterior  testis). 

The  ovary  is  situated  anteriorly  and  adjacent  to  the  foremost  testis,  usually  in 
the  median  line,  but  sometimes  moved  to  one  side  or  the  other  by  the  development 
of  the  uterus.  It  is  normally  wider  than  long,  oval,  with  smooth  margins  and 
measures  0-025-0-065  x  0-083-0-1  mm.  There  appears  to  be  a  seminal  receptacle 
lying  dorsally  to  the  ovary,  but,  despite  several  attempts,  it  has  not  been  possible  to 
obtain  satisfactory  serial  sections  of  the  female  complex,  so  whether  this  is  a  true 
seminal  receptacle  or  part  of  the  uterus  has  not  been  determined,  nor  has  the  course 
of  Laurer's  canal,  if  present.  The  viteilaria  consist  of  numerous  oval  follicles  in 
lateral  fields  reaching  from  about  the  posterior  border  of  the  ventral  sucker  to  near 
the  posterior  end  of  the  worm.  Behind  the  posterior  testis  the  fields  meet  in  the 
ventral  plane,  and  come  close  together  in  the  dorsal,  being  separated  only  by 
the  excretory  vesicle.  Between  the  testes,  and  between  the  anterior  testis  and  the 
ovary  the  follicles  may  extend  towards  the  median  line,  but  do  not  reach  it.  The 
vitelline  fields  are  sometimes  interrupted  opposite  the  ovary  and  the  testes.  The 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


161 


FIG.  4.     Bathycreadium  biscayense  sp.  nov.  :   specimen  from  Trachyrhynchus 
trachyrincus  (ventral  view). 


uterus  runs  from  the  ovary  to  the  posterior  margin  of  the  ventral  sucker,  and  con- 
tains a  few  (up  to  about  10)  large  eggs.     These  measure  80-92  /mi  x  43-52  /mi. 

MATERIAL.     British  Museum  (Nat.  Hist.)  Reg.  no.  1973.4.9.1-20  (syntypes). 

Bathycreadium  Kabata,  1961,  has  been  reduced  to  synonymy  with  Nicolla  Wisniew- 
ski,  1933,  by  Slusarski  (1971).  Kabata  did  not  compare  the  two  genera  so  it  was 
left  to  Pritchard  (1966)  to  list  the  differences  between  them.  Slusarski  dismisses 
Pritchard's  diagnostic  characters  by  saying  that  they  are  too  variable  in  Nicolla, 
and  he  is  probably  correct  in  this  assumption  in  respect  of  most  of  these  characters. 
It  does  seem,  however,  that  at  least  one  character  mentioned  by  Pritchard  may  be 
used  in  separating  these  two  genera,  along  with  another  feature  she  did  not  mention. 


162  R.    A.    BRAY 

As  she  noted,  it  appears  that  in  all  the  species  of  Nicolla  (with  one  exception,  i.e. 
N.  elongata)  the  vitellaria  reach  anteriorly  to  the  ventral  sucker,  whilst  in  Bathy- 
creadium  they  do  not  reach  further  forward  than  the  posterior  margin  of  the  ventral 
sucker.  It  is  also  apparent  that  the  normal  situation  of  the  posterior  testis  in 
Nicolla  is  close  to,  or  contiguous  with,  the  cyclocoel  with  few  or  no  vitelline  follicles 
between  the  posterior  testis  and  the  intestinal  arch  (again  N.  elongata  is  an  exception). 
In  Bathycreadium  there  is  at  least  a  testis  diameter,  usually  considerably  more, 
between  the  posterior  testis  and  the  cyclocoel,  and  the  bulk  of  the  vitellaria  is  in 
this  region.  From  its  differentiating  characteristics  it  is  evident  that  Nicolla 
elongata  Maillard,  1970,  from  Onos  tricirratus  in  the  Gulf  of  Lion  should  be  assigned 
to  Bathycreadium  elongatum  (Maillard,  1970)  n.  comb. 

It  may  also  be  mentioned  here  that  the  concept  of  Bathycreadium  as  given  above 
is  close  to  that  of  Coitocaecum  Nicoll,  1915,  as  represented  by  Yamaguti  (1971).  He 
seems  to  be  in  error,  however,  in  stating  that  the  cirrus-sac  is  '  retort-shaped  contain- 
ing elongate  saccular  or  tubular  seminal  vesicle,  prostatic  complex  and  ejaculatory 
duct'  in  Coitocaecum.  The  type-species  of  this  latter  genus,  C.  gymnophallum 
Nicoll,  1915,  has,  according  to  the  original  author,  'no  true  cirrus-pouch',  while 
Crowcroft  (1951)  who  re-examined  Nicoll's  material  states  that  there  is  'a  small 
membranous  cirrus-sac  enclosing  a  short,  terminal  portion  of  the  male  duct'. 

B.  biscayense  differs  from  the  other  two  species  in  the  genus,  B.flexicollis  Kabata, 
1961,  and  B.  elongatum  (Maillard,  1970),  in  the  anterior  extent  of  the  vitellaria,  in 
the  proximity  of  the  testes  and  the  ovary  and  in  the  sucker-ratio.  It  also  appears 
that  it  normally  contains  fewer  eggs  than  the  other  species.  The  egg-size  of  B. 
biscayense  differs  greatly  from  that  of  B.  elongatum,  but  is  similar  to  that  of  B. 
flexicollis  as  corrected  by  Pritchard  (1966)  and  verified  by  the  author  from  paratype 
specimens  in  the  collections  of  the  British  Museum  (Natural  History). 


Family  LEPOCREADIIDAE  Nicoll,  1935 
Lepidapedon  rachion  (Cobbold,  1858)  Stafford,  1904 

(Fig-  5) 

Distomum  rachion  Cobbold,  1858. 
Lepodora  rachiaea  :   Odhner,  1905. 

Host  and  locality  :  intestine  of  Phycis  blennoides,  Stn.  17. 

Nine  contracted  specimens  were  present.  They  are  small  worms  o-66-i  mm  long 
and  0-27-0-44  mm  wide,  being  widest  behind  the  ventral  sucker,  and  tapering  to- 
wards the  anterior  end.  The  cuticle  bears  rows  of  closely-set  narrow  pointed  spines. 
The  oral  sucker  is  subterminal,  measuring  0-097-0-115  mm  in  diameter,  and  leads  into 
a  prepharynx  which,  when  apparent,  varies  in  length  up  to  44  /xm  and  is  usually, 
but  not  invariably,  longer  than  the  oesophagus,  which  may  reach  a  length  of  28  jtrni. 
The  phary nx  is  large  and  pyriform  0-09-0- 13  mm  x  0-066-0-106  mm.  The  intestinal 
bifurcation  occurs  close  to  the  ventral  sucker,  and  the  caeca,  which  reach  to  the 
posterior  end  of  the  body,  are  wide.  The  rounded  ventral  sucker  lies  in  the  anterior 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


163 


FIG.  5.     Lepidapedon  rachion  (Cobbold)  :   ventral  view. 


half  of  the  worm,  and  measures  0-08-0-12  mm  in  diameter,  giving  an  oral :  ventral 
sucker  ratio  of  i  :  0-75-1-08.  Only  in  the  largest  specimen  was  the  ventral  sucker 
slightly  larger  than  the  oral,  whilst  in  the  other  specimens  the  oral  sucker  was 
distinctly  larger  than  the  ventral. 

The  excretory  pore  is  terminal  and  leads  into  a  simple  elongate  excretory  vesicle, 
which  has  been  traced  as  far  forward  as  the  posterior  testis. 

The  genital  pore  occurs  closely  anterior  to  the  ventral  sucker  and  to  the  left  of  the 
median  line  ;  in  fact,  it  lies  ventrally  to  the  left  caecum.  The  muscular  cirrus-sac 
reaches  to  near  the  posterior  margin  of  the  ventral  sucker,  measures  about  0-09  mm 
in  length  and  contains  a  coiled,  tubular,  seminal  vesicle  and  a  pars  prostatica,  which 
is  slightly  longer  than  the  short  muscular  cirrus.  The  external  seminal  vesicle  is 


164  R.    A.    BRAY 

also  tubular  and  is  coiled  amidst  a  mass  of  gland-cells.  This  mass  of  cells  extends 
behind  the  ventral  sucker  and  is  surrounded  by  a  thin  membrane.  In  the  posterior 
third  of  the  body  lie  the  testes,  in  tandem  formation.  They  are  transversely  elongate, 
have  smooth  or  indented  margins  and  measure  0-1-0-14 mm  x  0-03-0-05  mm 
(anterior)  and  0-08-0-14  mm  x  0-04-0-06  mm  (posterior). 

The  ovary  lies  just  anteriorly  to  the  foremost  testis  in  the  median  line  and  is 
separated  from  the  ventral  sucker  by  the  uterine  coils.  It  is  transversely  elongate, 
measuring  0-08-0-13  mm  x  0-04-0-07  mm,  with  smooth  or  slightly  indented  mar- 
gins. Dorsal  to  the  ovary  lies  a  globular  receptaculum  seminis.  The  vitelline  glands 
are  numerous  and  reach  anteriorly  as  far  as  about  the  middle  level  of  the  ventral 
sucker.  They  lie  laterally  and  ventrally  to  the  intestinal  caeca  and  reach  inwardly 
to  the  gonads,  and  although  they  intrude  between  the  gonads  they  do  not  meet  in 
the  median  line.  Posteriorly  to  the  testes,  however,  the  vitelline  fields  are  confluent 
in  the  median  line.  The  uterine  coils  lie  between  the  ovary  and  the  ventral  sucker 
and  contain  eggs,  all  of  which,  in  the  present  specimens,  are  collapsed.  In  this 
condition  they  measure  about  57-60  /mi  in  length. 

These  worms  are  recognizable  as  Lepidapedon  rachion  despite  the  fact  that,  pre- 
sumably due  to  contraction,  the  worms  are  on  average  shorter  than  is  normal  for 
this  species.  This  parasite  is  fairly  common  in  Gadidae  in  the  northern  Atlantic, 
but  has  not,  apparently,  been  recorded  from  this  host  before. 


Lepidapedon  elongatum  (Lebour,  1908)  Nicoll,  1910 

(Fig.  6) 
Lepodora  elongatum  Lebour,  1908. 

Hosts  and  localities  :  intestine  of  Lepidion  eques,  Stn.  i  ;  intestine  of  Trachyrhynchus 
trachyrincus,  Stn.  i,  Stri.  7,  Stn.  24. 

These  worms  were  common  in  the  intestine  of  both  hosts.  Their  normal  shape 
appears  to  be  elongate  oval,  but  some  of  the  present  specimens  are  contracted  to  a 
more  pyriform  shape,  whilst  some,  taken  from  a  frozen  specimen  of  L.  eques,  are 
very  elongate  in  the  forebody.  The  length  of  the  worm  varies  between  0-73  mm  and 
2-1  mm,  and  the  width  between  0-17  mm  and  0-42  mm.  The  cuticle  bears  rows  of 
narrow  pointed  spines.  Rounded  and  subterminal,  the  oral  sucker  measures 
0-09-0-17  mm  in  diameter.  The  ventral  sucker,  which  is  situated  in  the  middle 
third  of  the  body,  is  also  rounded  and  measures  0-06-0-13  mm  m  diameter.  The 
oral :  ventral  sucker  ratio  is  i  :  0-55-1.  The  prepharynx  in  grossly  contracted 
specimens  is  not  apparent,  but  in  less  contracted  specimens  it  varies  greatly  in  length, 
measuring  up  to  0-6  mm  in  one  specimen  from  a  frozen  L.  eques.  Normally,  the 
variation  appears  to  be  between  about  o-oi  mm  and  o-i  mm.  The  pharynx  is  longer 
than  wide,  measuring  0-06-0-14  mm  x  0-04-0-12  mm  and  has  a  tendency  to  be  of  a 
similar  size  to  or  somewhat  larger  than  the  ventral  sucker.  It  leads  into  the  oeso- 
phagus which,  like  the  prepharynx,  varies  considerably  in  length,  and  is  usually 
shorter  than  the  prepharynx,  being  0-0-06  mm  in  length.  It  bifurcates  about  midway 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


165 


FIG.  6.     Lepidapedon  elongatum  (Labour)  :   three  specimens  from  Trachyrhynchus 

trachyrincus  (ventral  view). 


between  the  suckers  or  nearer  to  the  ventral  sucker.  The  caeca,  lined  with  a  deeply 
staining  epithelium,  reach  to  near  the  posterior  end  of  the  body. 

The  excretory  pore  is  terminal  and  leads  into  a  simple  tubular  vesicle  which  has 
been  traced  anteriorly  to  the  level  of  the  hinder  margin  of  the  hinder  testis. 

The  genital  pore  is  situated  to  the  left  of  the  median  line  close  to  the  anterior 
margin  of  the  ventral  sucker  or  more  further  forward.  The  strongly  muscular,  oval 
cirrus-sac  measures  about  0-07-0-14  mm  x  0-04-0-09  mm,  but  is  not  easy  to  measure 
satisfactorily  due  to  the  angle  at  which  it  lies  to  the  body  surface.  It  contains  a 
coiled  tubular  seminal  vesicle,  a  distinctly  widened  pars  prostatica  and  a  muscular 
cirrus.  The  internal  seminal  vesicle  leads,  via  a  narrow  canal,  into  a  convoluted 


166  R.   A.    BRAY 

external  seminal  vesicle  surrounded  by  gland-cells.  These  cells  are  in  turn  surrounded 
by  a  very  thin  membrane,  and  reach  to  a  position  well  posterior  to  the  ventral  sucker. 
The  testes  have  smooth  margins  and  are  either  transversely  elongate  or  rounded. 
They  are  arranged  in  tandem  at  about  the  middle  of  the  hindbody.  The  anterior 
testis  measures  0-08-0-22  mm  x  0-04-0-13  mm  and  the  posterior  0-09-0-2  mm  x 
0-04-0-16  mm.  There  is  a  narrow  space  between  the  testes,  and  also  between  the 
anterior  testis  and  the  ovary,  which  lies  in  the  median  line,  and  is  separated  from  the 
ventral  sucker  by  the  uterine  coils.  Like  the  testes  it  may  be  either  transversely 
elongate  or  rounded,  and  measures  0-06-0-17  mm  x  0-04-0-11  mm.  The  seminal 
receptacle  lies  dorsally  to  the  ovary  and  Laurer's  canal  runs  to  the  dorsal  surface  at 
about  the  same  level.  The  vitelline  glands  do  not  reach  anteriorly  as  far  as  the 
ventral  sucker,  but  terminate  on  a  level  with  the  hinder  end  of  the  external  seminal 
vesicle,  or  thereabouts.  The  follicles  lie  mostly  in  a  plane  ventral  to  the  intestinal 
caeca,  with  some  lateral  to  them.  The  lateral  vitelline  fields  may  be  interrupted 
beside  the  ovary  and  testes  and  may  reach  to  the  median  line  between  these  organs, 
and  also  ventrally  to  the  uterus.  Behind  the  testes  the  fields  usually  merge  in  the 
median  line,  but  sometimes  the  excretory  vesicle  appears  to  form  a  barrier  between 
them.  The  uterus  runs  between  the  ovary  and  the  ventral  sucker,  and  opens  into 
the  genital  atrium  through  a  muscular  metraterm  of  about  the  same  length  as  the 
cirrus-sac.  The  uterus  contains  numerous  eggs,  the  few  uncollapsed  ones  measuring 
58-65  />im  long  by  about  30  /u,m  wide. 

Several  authors  have  used  the  name  Lepidapedon  elongatum  for  specimens  de- 
scribed from  various  species  of  fishes  from  the  west  and  east  coasts  of  North  America 
(Manter  1926  ;  Linton  1940  ;  Ching  1961),  the  Pacific  coast  of  Panama  (Caballero  y 
C.  el  al.  1955),  Bombay  (Gupta  and  Sehgal  1971),  the  Black  Sea  (Skrjabin  and  Koval 
1960)  and  the  Barents  Sea  (Dogiel  1936),  in  addition  to  those  originally  described 
from  Gadus  morhua  on  the  Northumberland  coast  of  England  by  Lebour  (1908). 
The  variation  represented  in  these  descriptions  indicates  that  more  than  one  species 
may  be  involved.  The  position  is  further  complicated  by  the  descriptions  of  very 
similar  trematodes  under  other  names,  for  example,  Lepidapedon  coelorhynchi 
Yamaguti,  1938,  L.  gadi  (Yamaguti,  1934)  and  L.  microcotyleum  Dollfus,  1953.  It 
seems  that  the  latter  two  species  may  be  distinguished  from  Lebour's  form  by  egg- 
size,  although  their  egg-size  overlaps  some  of  the  later  descriptions  of  '  L.  elongatum '. 
L.  coelorhynchi  was  considered  a  synonym  of  L.  elongatum  by  Hanson  (1950)  and  this 
has  been  accepted  by  most  subsequent  authors. 

The  specimens  described  above  differ  from  Lebour's  description  in  that  most  of 
them  have  a  ventral  sucker  distinctly  smaller  than  the  oral.  She  gives  measure- 
ments of  only  one  specimen,  the  ventral  sucker  being  0-12  mm  across  and  the  oral, 
o-i  mm  across.  It  should  be  noted,  however,  that  in  her  figure  the  ventral  sucker 
appears  to  be  slightly  smaller  than  the  oral.  The  distribution  of  the  vitellaria 
between  the  ovary  and  the  ventral  sucker  is  rather  variable  and  probably  not  a  good 
feature  to  use  for  differentiating  species  in  this  group,  as  was  suggested  by  Polyanski 
(1955).  Figure  6  illustrates  variation  in  three  specimens  from  the  intestine  of  the 
same  specimen  of  T.  trachyrincus,  but  at  the  same  time  they  also  show  a  certain 
constancy  of  morphological  features. 


FISH   TREMATODES   FROM   BAY    OF   BISCAY  167 

Family  AZYGIIDAE  Odhner,  1911 
Otodistomum  veliporum  (Creplin,  1837)  Stafford,  1904 

Host  and  locality  :  intestine  of  Dalatias  licha,  Stn.  i. 

This  trematode  has  been  reported  from  this  host  by  a  number  of  authors,  including 
Dollfus  (1937)  who  records  it  from  the  Bay  of  Biscay,  off  Biarritz,  and  also  gives  a 
list  of  previous  records.  Since  then  it  has  been  found  in  this  host  in  Norwegian 
waters  by  Braten  (1964)  and  in  the  Cook  Strait,  New  Zealand,  by  Manter  (1954). 

Family  ZOOGONIDAE  Odhner,  1911 
Steganoderma  abyssorum  (Odhner,  1911)  Manter,  1947 

(Fig-  7) 

Proctophantastes  abyssorum  Odhner,  1911. 
Deretrema  abyssorum  :   Price,  1934. 

Hosts  and  localities  :   intestine  of  Trachyrhynchus  trachyrincus ,  Stn.  i,  Stn.  7,  Stn. 

24  ;  intestine  of  Coelorinchus,  caelorhinchus ,  Stn.  i,  Stn.  37  ;  intestine  of  Lepidion 

eques,  Stn.  i. 

The  degree  of  contraction  of  the  fixed  worms  varies  considerably  so  that  the  shape 
varies  from  a  very  elongate  pyriform  to  a  short,  broadly  pyriform.  The  length  is 
between  0-62  mm  and  1-66  mm  and  the  greatest  width  is  0-28-0-63  mm,  which  occurs 
invariably  at  the  region  of  the  ventral  sucker.  The  rounded  oral  sucker  is  subter- 
minal,  0-18-0-32  mm  in  diameter,  whilst  the  ventral  sucker  is  considerably  larger, 
situated  somewhat  posteriorly  to  the  middle  of  the  worm,  and  is  wider  than  long, 
being  0-25-0-6  mm  across  by  0-18-0-44  mm-  Transversely  across  the  inside  of  the 
ventral  sucker  runs  a  low  muscular  ridge.  The  oral :  ventral  sucker  width  ratio  is 
i  :  1-08-2-1  and  the  sucker  length  ratio  is  i  :  0-9-1-6.  Only  in  extended  specimens 
is  a  prepharynx  to  be  seen,  and  it  reaches  to  no  greater  length  than  0-04  mm.  The 
small  globular  pharynx  has  a  diameter  of  0-03-0-08  mm  and  from  it  leads  an  oeso- 
phagus which  varies  greatly  in  length  between  0-08-0-22  mm.  In  grossly  contracted 
specimens  it  has  not  been  observed.  It  bifurcates  about  halfway  between  the 
suckers  and  the  caeca  are  fairly  short,  terminating  dorsally  to  the  testes  or  vitellaria. 

The  excretory  pore  is  terminal  and  leads  into  a  vesicle  which  reaches  forward, 
dorsally  to  the  uterus,  for  an  undetermined  distance  and  as  far  as  can  be  seen  is  a 
simple  tube. 

Approximately  on  a  level  with  the  intestinal  bifurcation,  in  a  submarginal  position, 
lies  the  genital  pore.  The  cirrus-sac  reaches  past  the  anterior  margin  of  the  ventral 
sucker,  and  in  contracted  specimens  nearly  reaches  to  the  transverse  ridge,  measuring 
0-2-0-37  mm  x  0-06-0-09  mm-  It  contains  a  coiled  seminal  vesicle,  the  coils  of 
which  are  often  tightly  packed,  a  well-developed  pars  prostatica  invested  with  a  mass 
of  gland  cells  and  a  cirrus,  of  about  one-third  the  length  of  the  cirrus-sac.  The 
proximal  half  of  the  cirrus  appears  to  be  lined  with  a  cuticle  which  may  be  thrown 
in  wrinkles.  The  two  oval  or  elongate  oval  testes  are  always  partly  obscured  by  the 


1 68 


R.    A.    BRAY 


FIG.  7.     Steganoderma  abyssorum  (Odhner)  :   two  specimens  from  Trachyrhynchus 

trachyrincus  (ventral  view). 


eggs  or  the  vitelline  glands  and  lie  symmetrically  one  on  each  side  of  the  body, 
separated  from  the  posterior  border  of  the  ventral  sucker  by  some  of  the  vitellaria. 
They  measure  about  0-16-0-22  mm  x  0-13-0-2  mm. 

The  rounded  ovary  is  smaller  than  the  testes,  being  about  0-15  mm  in  diameter. 
It  is  situated  more  or  less  on  the  posterior  border  of  the  ventral  sucker,  but  sometimes 
entirely  posterior  or  anterior  to  it.  Posteriorly  and  dorsally  to  the  ovary  lies 
the  rounded  or  oval  seminal  receptacle,  which  is  rather  smaller  than  the  ovary. 
Immediately  posterior  to  the  ventral  sucker,  in  two  symmetrical  lateral  groups,  are 
the  vitellaria,  which  lie  somewhat  ventrally  to  the  testes,  and  overlap  them  in  vary- 
ing degrees.  There  are  between  five  and  ten  follicles  in  each  group.  The  uterus  fills 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  169 

most  of  the  area  posterior  to  the  ventral  sucker,  and  therefore  in  the  majority  of 
specimens  the  details  of  the  reproductive  organs  are  obscured.  It  contains  numerous 
operculate  eggs  measuring  34-41  />tm  long  by  16-21  /mi.  The  metraterm  is  long, 
muscular  and  helical.  It  enters  a  small,  muscular  genital  atrium,  which  is  surrounded 
by  a  scattering  of  glandular  cells. 

This  species  was  originally  described  as  the  type-species  of  the  genus  Procto- 
phantastes  Odhner,  1911,  from  Macrurus  (Coryphaenoides)  rupestris  and  Gadus 
aeglefinus  from  Norway.  Price  (1934)  considered  Proctophantastes  to  be  a  synonym 
of  Deretrema  Linton,  1910,  but  this  synonymy  was  questioned  by  Manter  (1947) 
who  considered  it  a  synonym  of  Steganoderma  Stafford,  1904.  Odhner  (191  ia)  does 
not  discuss  the  relationship  of  Proctophantastes  with  either  of  the  genera  with  which 
it  has  been  synonymized.  Whilst  Price  (1934)  makes  his  synonymy  with  little 
comment,  Manter  (1947)  discusses  the  relationship  of  the  three  genera  in  some  detail. 
As  has  been  mentioned  above,  the  latter  author  considers  Proctophantastes  to  be  a 
synonym  of  Steganoderma  and  he  suggests  that  the  differences  between  the  type- 
species  of  Steganoderma,  S.  formosum  Stafford,  1904,  and  P.  abyssorum  are  of  specific 
rather  than  generic  value.  These  differences  are  in  sucker-size,  oesophagus-length 
and  length  of  vitelline  fields.  The  present  author  agrees  with  Manter's  conclusion. 
Therefore,  the  differences  Manter  (1947)  indicates  between  Steganoderma  and  Dere- 
trema also  apply  to  Proctophantastes  when  compared  with  Deretrema.  In  short, 
Price's  synonymy  is  rejected,  because  the  vitellaria  in  Deretrema  do  not  reach  pos- 
teriorly to  the  ventral  sucker,  whilst  in  Steganoderma  and  Proctophantastes  the  majority 
of  the  vitellaria  are  situated  in  this  region.  Subsequent  to  Manter  (1947),  only 
Yamaguti  (1953,  1958,  1970,  1971)  has  continued  to  consider  Proctophantastes  a 
valid  genus.  The  features  he  uses  to  distinguish  Proctophantastes  from  Steganoderma 
are,  I  believe,  not  sufficient  to  uphold  the  genus.  In  1953  he  indicates  that  in 
Proctophantastes  the  vitellaria  are  compact,  while  in  Steganoderma  they  are  in  longi- 
tudinally elongate  groups.  This  type  of  difference  would  appear  to  be  of  specific 
rather  than  generic  importance,  as  it  concerns  fairly  small  details  of  the  vitelline 
extent.  Yamaguti  (1958)  distinguishes  the  genera  on  the  shape  of  the  cirrus-sac 
and  the  position  of  the  ventral  sucker.  An  examination  of  Miller's  (1941)  redescrip- 
tion  of  Stafford's  specimen  of  S.  formosum,  and  also  of  the  other  descriptions  of  this 
species  given  by  Manter  (1925,  1926),  Linton  (1940),  Ching  (1960),  Ronald  (1960) 
and  Strelkov  (1960)  shows  that  the  cirrus-sac  of  this  species  resembles  that  of  P. 
abyssorum  in  shape.  The  ventral  sucker  in  this  latter  species  lies  just  behind  the 
middle  of  the  body,  while  in  S.  formosum  it  lies  just  anterior  to  the  centre  of  the  body. 
This  condition  probably  varies  with  the  contraction  or  growth  of  the  specimen,  so 
that  its  value  as  a  generic  character  is  rather  doubtful.  In  1970,  Yamaguti  suggested 
that  a  'profusely  developed  "periatrial  gland"  may  be  a  generic,  not  a  specific 
character,  though  not  mentioned  by  Odhner '.  He  is  referring  to  a  character  present 
in  a  new  species  he  is  describing,  and  this  character  is,  I  believe,  a  feature  of  Neo- 
steganoderma,  and  is  discussed  in  connection  with  the  following  species.  The  few 
scattered  gland-cells  around  the  genital  atrium  in  the  present  specimen  do  not,  it 
appears,  represent  a  feature  of  generic  importance,  and,  moreover,  similar  glands 
have  been  described  in  other  species  of  Steganoderma,  e.g.  S.  elongatum  Manter,  1947. 


iyo  R.   A.    BRAY 

Finally,  Yamaguti  (1971)  upholds  Proctophantastes  on  account  of  the  characteristic 
structure  of  the  genital  atrium.  There  is,  however,  no  peculiar  feature  in  the  genital 
atrium  of  the  present  specimens,  nor  is  it  indicated  in  Odhner's  generic  definition 
where  he  merely  says  '  Genitalsinus  ziemlich  klein'.  As  the  various  attempts  to 
validate  Proctophantastes  do  not  appear  convincing,  this  genus  is  here  considered  to 
be  a  synonym  of  Steganoderma. 

The  present  specimens  show  no  outstanding  morphological  differences  from  P. 
abyssorum,  as  described  by  Odhner  (igiia),  and  are  therefore  considered  to  belong 
to  this  species.  There  are,  however,  two  minor  differences,  the  first  being  the  sparse 
gland-cells  around  the  genital  atrium  and  the  second  the  presence  of  a  ridge  running 
transversely  across  the  inner  surface  of  the  ventral  sucker.  This  latter  feature  was 
not  easy  to  detect  in  many  whole  mounts  and  may  easily  have  been  overlooked  by 
Odhner. 


Neosteganoderma  glandulosum  Byrd,  1964,  emend 

(Fig.  8) 

Proctophantastes  glandulosa  :   Yamaguti,  1971. 

Host  and  locality  :   intestine  of  Beryx  decadactylus ,  Stn.  37. 

All  the  worms  are  contracted  to  a  broadly  pyriform  shape,  which  tapers  towards 
the  anterior  end.  The  length  varies  between  i  mm  and  1-38  mm  and  the  maximum 
width,  which  occurs  at  the  level  of  the  testes,  between  0-76  mm  and  0-96  mm. 
Anteriorly,  the  cuticle  bears  long,  narrow  spines  which  also  occur  near  the  posterior 
extremity.  The  subterminal  oral  sucker  is  rounded,  0-27-0-38  mm  across  by 
0-26-0-33  mm  long.  At  about  the  middle  of  the  worm,  or  just  behind,  lies  the 
transversely  elongate  ventral  sucker,  measuring  0-69-0-77  mm  across  by  0-27-0-33 
mm  long,  reaching  close  to  the  lateral  margins  of  the  body  on  either  side.  The 
oral :  ventral  sucker  width  ratio  is  i  :  2-1-2-8,  whilst  the  length  of  both  suckers  is 
almost  the  same.  A  small  prepharynx  can  be  seen  in  serial  section.  It  enters  a 
globular  pharynx  of  about  0-11-0-13  mm  diameter.  The  oesophagus  is  fairly  short, 
curved  dorsally  to  the  pharynx,  and  bifurcates  just  posteriorly  to  the  pharynx.  The 
caeca  are  quite  short,  reaching  to  a  position  dorsal  to  the  testes. 

The  excretory  pore  is  terminally  situated,  or,  when  the  worm  is  greatly  distended 
with  eggs,  it  may  appear  to  be  slightly  dorsal.  The  saccular  vesicle  reaches  as  far 
as  the  posterior  border  of  the  ovary,  and  in  this  region  part  of  it  lies  ventrally  to  the 
seminal  receptacle. 

On  the  left  side  of  the  body,  submarginally  and  just  anterior  to  the  ventral  sucker 
lies  the  genital  pore.  The  genital  atrium  has  a  strong  muscular  wall,  with  irregular 
outpocketings.  Into  it  leads  a  large  cirrus-sac,  the  bulk  of  which  lies  just  anterior 
to  the  ventral  sucker  and  at  right  angles  to  the  long  axis  of  the  worm.  The  cirrus-sac 
reaches  transversely  across  the  body  to  the  right  intestinal  caecum,  where  it  twists 
and  runs  slightly  posteriorly  towards  the  centre  of  the  ventral  sucker,  in  a  plane  dorsal 
to  the  remainder  of  the  cirrus-sac.  The  relatively  small,  coiled  internal  seminal 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


171 


I72  R.    A.    BRAY 

vesicle  leads  into  a  well-developed  pars  prostatica,  which,  with  its  innumerable  as- 
sociated glands,  fills  most  of  the  cirrus-sac.  The  muscular  cirrus  is  shorter,  contain- 
ing some  long,  narrow  spines,  lying  lengthwise,  but  as  they  have  been  seen  clearly 
only  in  serial  section  the  overall  shape  of  these  spines  has  not  been  made  out.  The 
testes  are  arranged  symmetrically,  just  posterior  to  the  ventral  sucker  and  close  to 
the  lateral  margins  of  the  worm.  Apparently,  normally  they  are  of  an  oval  shape, 
but  with  growth  of  the  uterus  certain  lobe-like  processes  appear.  In  one  specimen 
the  testes  measured  0-27  x  0-3  mm  and  0-4  x  0-19  mm. 

Between  the  testes,  and  lying  towards  the  right  of  the  worm,  is  the  ovary.  In 
one  specimen  it  was  seen  to  be  of  an  irregular  quadrilateral  shape,  0-27  x  0-27  mm. 
The  large,  oval  receptaculum  seminis  lies  dorsally  to  the  ovary  and  the  left  testis. 
It  may  measure  0-3  x  0-19  mm.  Ventral  to  the  anterior  extremity  of  the  recepta- 
culum seminis  is  a  well-developed  'shell '-gland.  The  vitelline  follicles  are  situated 
in  two  lateral  fields,  ventrally  to  the  testes  and  reaching  to  about  the  posterior  margin 
of  the  ventral  sucker.  Each  field  contains  four  to  eight  follicles.  The  egg-filled 
uterus  occupies  the  great  majority  of  the  hindbody,  obscuring  the  reproductive 
organs  in  this  region.  The  metraterm  is  long  and,  particularly  near  the  genital 
atrium,  strongly  muscular.  It  enters  the  latter  from  the  dorsal  side.  For  most  of 
its  length  it  is  surrounded  by  an  investment  of  gland-cells  (Fig.  8b).  Where  the 
metraterm  and  the  cirrus-sac  run  adjacent  to  each  other  the  cells  invest  both  organs 
and  also  surround  the  genital  atrium.  The  operculate  eggs  measure  26-31  jum  x 
16-21  /u,m. 

Byrd  (1964)  described  Neosteganoderma  glandulosum  from  a  beryciform  fish, 
Polymixia  lowei,  in  the  Straits  of  Florida.  He  differentiated  his  new  genus  from  other 
zoogonids  with  follicular  vitellaria  situated  posteriorly  to  the  ventral  sucker  by  a 
number  of  features.  These  included  the  posterior  position  of  the  ventral  sucker  and 
the  elongate  oesophagus  and  forebody,  but  as  can  be  seen  from  the  present  contracted 
specimens  these  are  not  reliable  features.  It  should  be  noted  that  Byrd's  specimens 
were  treated  with  chloretone  in  sea-water  before  fixation,  which  probably  accounted 
for  the  relaxation  of  the  forebody.  The  other  distinguishing  characteristics  he 
mentions  are  the  very  large  ventral  sucker  and  the  '  large  glandular  mass  surrounding 
the  genital  pore'.  The  latter  feature  is  very  prominent  in  the  present  specimens 
and  appears  to  be  the  most  reliable  feature  for  distinguishing  Neosteganoderma  from 
the  closely-related  genus  Steganoderma.  Yamaguti  (1971),  however,  places  Byrd's 
species  in  Proctophantastes ,  which,  as  pointed  out  above,  does  not  possess  a  large 
mass  of  gland-cells  surrounding  the  genital  atrium.  It  appears  that  Yamaguti  is 
basing  his  definition  of  Proctophantastes  Odhner,  1911,  not  on  its  type-species,  P. 
abyssorum  Odhner,  1911,  but  on  his  own  species  P.  polymixiae  Yamaguti,  1970,  from 
Polymixia  japonica  in  the  Pacific  Ocean  off  Hawaii,  which  he  describes  as  having  a 
large  'periatrial  gland'.  This  suggests  that  this  species  should  become  Neostegano- 
derma polymixiae  (Yamaguti,  1970)  n.  comb.  It  seems  possible  that  it  may  be 
found  to  be  conspecific  with  N.  glandulosum. 

As  stated  above,  certain  features  of  the  morphology  of  the  present  specimens  are 
different  from  those  described  by  Byrd,  but  these  are  considered  to  be  due  to  the 
differences  in  the  fixation  technique  used.  The  oesophagus  in  the  present  forms  is 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  173 

particularly  reduced  by  comparison  with  the  original  description.  Byrd's  statement 
that  the  metraterm  enters  the  genital  atrium  from  the  ventral  side  appears  to  be  a 
typographical  error,  as  a  close  examination  of  his  figures  shows  that  in  his  specimens, 
as  in  mine,  the  metraterm  enters  the  genital  atrium  from  the  dorsal  side. 


Family  FELLODISTOMIDAE  Nicoll,  1913 
(?)  Steringotrema  diver  gens  (Rud,  1809)  Odhner,  1911 

(Fig-  9) 

Distoma  diver  gens  Rud.,  1809. 
Fasciola  diver  gens  :   de  Blainville,  1820. 

Host  and  locality  :  intestine  of  Capros  aper,  Stn.  38,  Stn.  53. 

Seven  specimens  were  recovered  from  a  fish  at  Stn.  38  and  two  from  a  fish  at  Stn. 
53.  These  specimens  are,  unfortunately,  rather  contracted,  but  enough  detail  has 
been  observed  to  suggest  that  they  probably  belong  to  Steringotrema  diver  gens. 

The  worms  are  oval,  often  tapering  at  each  end,  and  measure  0-75-1-2  mm  long 
by  0-42-0-47  mm  in  maximum  width  at  the  level  of  the  ventral  sucker.  The  cuticle 
is  unarmed.  The  subterminal  oral  sucker  is  0-11-0-15  mm  m  diameter  and  leads 
into  a  globular  pharynx  of  0-09-0-11  mm  diameter.  Because  of  the  contraction 
the  oesophagus  is  curved,  but  it  bifurcates  just  about  the  level  of  the  genital  pore, 
and  the  intestinal  caeca  extend  to  about  the  level  of  the  testes.  About  halfway 
along  the  body  lies  the  transversely-elongate  ventral  sucker,  measuring  0-21- 
0-31  mm  x  0-38-0-42  mm,  giving  an  oral :  ventral  sucker  ratio  of  about  i  :  2-7-3-6. 

The  excretory  pore  is  terminal,  and  the  vesicle  is  V-shaped,  the  limbs  of  which 
have  been  traced  almost  to  the  ventral  sucker  in  serial  sections. 

About  halfway  between  the  suckers  and  just  to  the  left  of  the  median  line  lies  the 
genital  pore.  The  rather  stout  cirrus-sac,  0-18-0-19  mm  long  and  0-11-0-13  mm 
wide,  contains  a  bipartite  seminal  vesicle  leading  into  an  elongate  pars  prostatica 
which  passes  from  the  anterior  end  of  the  seminal  vesicle  to  near  the  posterior  end 
of  the  cirrus-sac  before  turning  and  running  anteriorly  to  unite  with  a  short,  muscular, 
smooth  cirrus.  Due  to  contraction  of  the  body,  the  testes,  the  ovary  and  the  egg- 
filled  uterus  are  pressed  closely  against  the  ventral  sucker.  Thus,  as  the  eggs  are 
opaque,  the  testes  and  the  ovary  are  discernible  only  in  serial  section.  The  testes 
lie  symmetrically  at  the  lateral  margins  of  the  body,  with  the  ovary  lying  close  to  the 
right  testis.  The  vitelline  follicles  are  arranged  in  four  groups,  two  on  each  side  of 
the  worm,  one  group  anterior  to  the  ventral  sucker  and  the  other  group  posterior  to 
the  sucker.  The  uterus  fills  almost  the  whole  of  the  hindbody  of  the  worm  and 
contains  numerous  operculate  eggs,  46-48  /xm  x  33-37  ju.m  with  shell  about  2-3  /xm 
thick. 

Any  slight  morphological  differences  which  can  be  detected  between  the  above 
description  and  those  given  by  Odhner  (191  ib),  Palombi  (1931)  and  Mathias  (1934) 
may  probably  be  explained  by  the  contraction  of  the  present  worms.  The  eggs,  as 
measured  when  teased  from  the  body  as  well  as  in  utero,  are  rather  wider  than  those 


174 


R.    A.    BRAY 


FIG.  9.     (?)  Steringotrema  divergens  (Rud.)  :   diagrammatic  ventral  view  with  intestine 

omitted. 


previously  recorded.  A  more  definite  statement  on  the  identity  of  these  specimens 
awaits  better-preserved  material.  It  should  be  noticed  that  the  previously-reported 
hosts  of  S.  divergens  (i.e.  gobies  and  blennies)  have  a  rather  different  biology  from  that 
of  the  present  host. 

Steringophorus  blackeri  sp.  nov. 
(Fig.  10) 

Hosts  and  localities  :  intestine  and  body-cavity  of  Xenodermichthys  copei  (type-host), 
Stn.  24  ;  intestine  of  Xenodermichthys  sp.  indet.,  National  Institute  of  Oceano- 
graphy, Stn.  6429  -  28°07'  N.,  I3°52'  W.  depth  0-390  m. 

Four  whole  specimens  were  recovered  from  X.  copei.  One  whole  specimen  and 
an  anterior  fragment  were  found  in  the  body-cavity  of  the  same  fish  specimen,  the 
fish  itself  having  itself  been  preserved  in  formalin.  The  remainder  of  the  specimens 
from  X.  copei  and  the  two  specimens  found  in  X.  sp.  indet.  were  taken  from  the 
intestine.  The  worms  occurred  either  singly  or  in  pairs  in  their  hosts.  The  whole 
worm  from  the  body-cavity  (Fig.  lob)  was  of  a  shape  and  size  rather  different  from 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


175 


0-1mm 


I76 


R.   A.   BRAY 


those  in  the  intestine  (Fig.  loa),  two  of  which  were  sectioned  for  further  study.  The 
other  specimens,  collected  by  the  author  from  fishes  in  the  collection  of  the  National 
Institute  of  Oceanography,  included  one  which  was  apparently  young  (Fig.  ice)  and 
an  older  worm  (Fig.  loe).  Despite  the  variation  in  size,  the  internal  morphology 
indicates  clearly  that  all  these  specimens  represent  the  same  species. 

The  'intestinal  form'  from  X.  copei  is  elongate,  being  widest  just  posteriorly  to 
the  ventral  sucker,  which  is  about  one-fifth  of  the  body-length  from  the  anterior  end. 
The  forebody  tapers  sharply,  while  the  tapering  of  the  hindbody  is  less  pronounced, 
and  it  is  rounded  terminally.  The  forms  from  X.  sp.  indet  are  much  smaller,  but 
with  a  similarly  shaped  forebody,  and  are  widest  just  posteriorly  to  the  ventral 
sucker  (at  about  one-third  of  the  body-length  from  the  anterior  end) ,  the  hindbody 
having  slightly  tapering  lateral  margins  and  a  rather  truncate  posterior  end.  The 
'  body-cavity  form '  from  X.  copei  has  a  similar  forebody  to  the  other  forms,  but  the 
hindbody  is  considerably  swollen  by  the  large  bulbous  excretory  vesicle,  the  limbs 
of  which  reach  into  the  forebody,  but  do  not  appear  to  be  swollen  in  the  same  way. 

The  dimensions  of  the  various  forms  are  included  in  Table  2. 


TABLE  2 

Measurements  of  Steringophorus  blackeri  sp.  nov. 


Host 

X.  copei 

X.  copei 

X.  sp.  indet. 

X.  sp. 

indet. 

'Form' 

Intestinal 

Body-cavity 

Younger 

Older 

No.  of  specimens 

3  (2  sectioned) 

i  +  fragment 

i 

i 

Length  (mm) 

4-6-9 

2-7 

2-2 

3-08 

Width  (mm) 

2-03-2-04 

1-45 

0-8 

1-68 

Oral  sucker 

0-47 

0-34-0-35 

0-243 

0-4 

diameter  (mm) 

' 

Ventral  sucker 

°'43 

0-29-0-33 

0-240 

o-45 

diameter  (mm) 

Oral  :  ventral 

i  :  0-92 

i  :  0-85-0-94 

i  :  0-98 

i  :  1-12 

sucker  ratio 

Pharynx  (mm) 

0-32  x  0-26 

0-22-0-26  x 

0-125  x  o-ii 

0-21  x 

0-25 

Oesophagus  (mm) 
Cirrus-sac  (mm) 

Ovary  (mm) 
Testes  (mm) 

Eggs  (/*m) 


0-21-0-19 

0-13-0-23  0-14  0-105  0-16 

about  0-32-0-35  about  0-21-0-32  0-26  x  0-14  0-48  x  0-16 

x  0-19  x  0-12 

0-27  dia.  0-21  x  0-16  0-18  x  o-u  0-21  x  0-19 

about  0-21  x  o-n-o-12  x  0-07-0-13  x  0-13-0-14  x 

0-18  0-16  0-14  0-17-0-2 

57-63  x  32-34  52-58  x  27-30  54  x  27-30  54-57  x  28-30 


The  cuticle  bears  no  spines,  but  has  numerous  transverse  wrinkles.  The  oral  and 
ventral  suckers  are  rounded,  and  with  one  exception  the  oral  is  slightly  larger  than 
the  ventral.  In  some  specimens  from  X.  copei  contraction  has  caused  the  opening 
of  the  ventral  sucker  to  be  disposed  anteriorly.  There  is  no  prepharynx,  so  the 
almost  globular  pharynx  abuts  directly  with  the  oral  sucker.  It  leads  into  a  fairly 
short  oesophagus,  which  bifurcates  at  about  the  level  of  the  anterior  margin  of  the 
ventral  sucker,  or  slightly  further  forward.  The  intestinal  caeca  are  lined  with  a 


FISH   TREMATODES   FROM   BAY   OF   BISCAY  177 

glandular  epithelium  and  run  posteriorly  to  a  point  normally  amidst  the  uterine 
coils  where  they  terminate,  about  halfway  along  the  hindbody. 

The  excretory  pore  is  terminal  and  leads  into  a  very  small  globular  vesicle  (of 
about  0'03-o-o6  mm  in  diameter)  before  entering  the  main  Y-shaped  vesicle.  The 
stem  of  the  vesicle  reaches  to  about  the  middle  of  the  body,  whilst  the  limbs  reach 
anteriorly  to  the  oral  sucker.  The  stem  may  be  very  wide  or  narrow,  but  the  limbs 
are  constantly  narrow.  The  excretory  pore  itself  appears  to  be  on  a  short 
protuberance  at  the  base  of  a  depression  and  is  surrounded  by  glandular  cells. 

The  genital  pore  is  situated  on  a  level  with,  or  just  anterior  to,  the  intestinal  bifur- 
cation and  slightly  to  the  left  of  the  median  line.  It  leads  into  a  muscular  genital 
atrium.  The  cirrus-sac  overlaps  the  anterior  border  of  the  ventral  sucker  and  con- 
tains a  relatively  small,  bipartite  seminal  vesicle,  a  long,  coiled  pars  prostatica  and 
a  short  muscular  cirrus.  The  testes  lie  posteriorly  to  the  ventral  sucker  a  little 
obliquely  to  each  other,  with  the  left  testis  being  slightly  anterior  to  the  right.  They 
are  close  to  the  ovary  and  to  each  other,  are  small  by  comparison  with  the  body-size 
and  have  more  or  less  deeply  indented  margins.  The  multilobate  ovary  lies  anteriorly 
and  sometimes  slightly  obliquely  to  the  right  testis.  It  is  associated  with  a  large 
'  shell  '-gland,  about  the  same  size  as  the  ovary,  lying  between  it  and  the  left  testis. 
Laurer's  canal  opens  dorsally,  level  with  the  '  shell  '-gland,  and  may  contain  a  small 
amount  of  spermatozoa  in  its  proximal  region.  Sperm  may  also  lie  in  the  proximal 
part  of  the  uterus.  The  vitellaria  consists  of  numerous  globular  or  short  trans- 
versely-elongate follicles  disposed  in  two  symmetrically  placed  fields  near  the  lateral 
margins  of  the  body.  These  fields  reach  from  about  the  level  of  the  anterior  border 
of  the  ventral  sucker  to  a  position  some  distance  posterior  to  the  testes.  The  uterus 
runs  in  the  median  line  from  the  ovary  to  about  the  middle  of  the  worm,  in  a  series  of 
lateral  coils.  It  then  forms  a  descending  series  of  loops  on  each  side  of  the  body  to 
about  two-thirds  of  the  way  along  the  hindbody  before  passing  anteriorly  again  as  a 
series  of  loops.  It  has  not  been  possible  to  ascertain  whether  the  uterus  passes  down 
one  side  of  the  body  and  then  down  the  other,  or  whether  it  divides  before  passing 
down  both  sides.  The  later  coils  of  the  uterus  pass  forward  in  the  median  line 
ventrally  to  the  descending  coils  and  the  gonads.  In  the  specimens  with  the  swollen 
excretory  vesicle  the  lateral  uterine  slings  are  forced  to  lie  close  to  the  margins  of 
the  body.  Within  the  uterus  lie  numerous  eggs,  many  of  which  are  collapsed,  but 
which  appear  to  be  operculate,  with  a  brownish-yellow  shell.  This  shell  is  covered 
with  numerous  small  bosses  giving  the  surface  a  granular  appearance,  which  is 
rather  less  marked  in  the  young  specimen  from  X.  sp.  indet. 

MATERIAL.     British  Museum  (Natural  History)  Reg.  no.  1973.4.9.21-24  (syntypes). 

Although  some  recent  authors  have  followed  Yamaguti  (1953)  in  considering 
Steringophorus  Odhner,  1905,  a  synonym  of  Fellodistomum  Stafford,  1904,  it  would 
seem  that  there  may  be  one  feature  differentiating  these  genera,  if  the  distribution  of 
the  vitellaria  is  taken  into  consideration.  In  Fellodistomum  the  vitellaria  extend 
anteriorly  well  beyond  the  ventral  sucker,  whilst  in  Steringophorus  they  do  not.  A 
close  examination  of  material  of  the  type-species  of  both  genera  from  the  type-hosts 
may  well  indicate  other  differences,  particularly  in  the  length  of  the  caeca  and  the 
configuration  of  the  fully-developed  uterus. 


178  R.   A.    BRAY 

This  new  species  would  appear  to  be  most  closely  related  to  Steringophoms  magnus 
Manter,  1934,  from  an  unidentified  eel-like  fish  in  300  fathoms  at  Tortugas,  Florida. 
It  differs,  however,  in  the  granular  pattern  of  the  egg-shell,  in  the  sucker  ratio,  in  the 
distribution  of  the  vitellaria  and  in  the  configuration  of  the  uterus  in  the  hindbody. 
The  sculpturing  on  the  egg-shell  seems  to  be  similar  to  that  described  in  Benthotrema 
plenum  Manter,  1934,  and  it  may  be  noted  that  although  Manter  (1934)  originally 
considered  this  to  be  a  generic  character,  later  (1954)  he  described  a  further  species 
in  the  genus,  B.  richardsoni,  in  which  the  eggs  apparently  have  smooth  shells. 

Family  HEMIURIDAE  Liihe,  1901 
Hemiurus  appendiculatus  (Rud.,  1802)  Looss,  1899 
Host  and  locality  :  stomach  of  Alosa  fallax,  Stn.  41. 

Seven  specimens  were  found.  This  is  a  common  parasite  of  shads  and  has  been 
recorded  in  A .  fallax  [A .  finta]  in  various  localities  in  the  Mediterranean  and  north- 
eastern Atlantic.  Nevertheless,  H.  appendiculatus  of  de  Oliveira  Rodrigues  et  al. 
(1972)  recorded  from  A .  fallax  from  the  coast  of  Portugal  may  not  belong  to  this 
species,  as  it  has,  judging  from  the  authors'  figure,  a  relatively  short  pars  prostatica 
and  an  undivided  seminal  vesicle.  The  normal  condition  appears  to  be  that  rep- 
resented by  the  present  specimens,  in  which  the  pars  prostatica  is  convoluted  pos- 
teriorly to  the  ventral  sucker,  and  in  which  the  anterior  part  of  the  bipartite  seminal 
vesicle  has  a  thick  muscular  wall. 

Lecithocladium  excisum  (Rud.,  1819)  Liihe,  1901 
Host  and  locality  :   stomach  of  Scomber  scombrus,  Stn.  44. 

Lecithochirium  gravidum  Looss,  1907 
Host  and  locality  :   stomach  of  Conger  conger,  Stn.  7. 

In  their  revision  of  Lecithochirium  Liihe,  1901,  Nasir  and  Diaz  (1971)  consider 
L.  gravidum  to  be  a  synonym  of  L.  rufoviride  (Rud.,  1819).  These  two  species  are 
certainly  very  similar  morphologically  and  according  to  Looss  (1908)  and  Guiart 
(1938)  they  are  distinguishable  only  by  size  and  sucker  ratio.  L.  gravidum  is  small 
with  an  oral :  ventral  sucker  ratio  of  about  i  :  2  whilst  rufoviride  is  larger  and  has 
suckers  of  more  or  less  equal  size.  The  present  specimens  (six  in  number)  have  an 
oral  sucker  diameter  of  0-24-0-29  mm  and  a  ventral  sucker  diameter  of  0-47-0-52  mm, 
and  therefore  a  sucker  ratio  of  i  :  1-8-2-1.  In  an  attempt  to  discover  whether  or 
not  the  sucker  ratio  in  this  group  is  influenced  by  allometric  growth  60  specimens, 
assigned  to  one  or  other  of  these  species,  were  measured.  These  specimens,  from  the 
collections  of  the  British  Museum  (Natural  History),  were  all  from  Conger  conger 
collected  at  Plymouth  and  Guernsey.  The  sucker  ratio  was  then  plotted  against  the 
body-length  as  is  shown  in  Fig.  n.  As  can  be  seen  there  are  two  distinct  regions, 
one  representing  those  specimens  with  a  sucker  ratio  of  more  than  1:1-7  and  the 
other  representing  those  with  a  ratio  of  less  than  i  :  1-6.  Allometric  growth  does 


FISH   TREMATODES   FROM   BAY   OF   BISCAY 


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FISH  TREMATODES   FROM   BAY   OF   BISCAY  181 

not  appear  to  have  a  significant  effect  on  the  sucker  ratio.  It  can  therefore  be  seen 
that,  commonly  occurring  in  Conger  conger,  there  are  two  very  similar  species 
differing  in  their  sucker  ratio,  if  not  in  any  other  obvious  feature.  This  informa- 
tion seems  to  uphold  the  validity  of  the  species  L.  gravidum,  to  which  the  present 
specimens  are  assigned. 

Derogenes  various  (Miiller,  1784)  Looss,  1901 

Hosts  and  localities  :  stomach  of  Molva  macrophthalma,  Stn.  43  ;  stomach  of 
Trachyrhynchus  trachyrincus ,  Stn.  43  ;  stomach  of  Micromesistius  poutassou, 
Stn.  6  ;  stomach  of  Phycis  blennoides,  Stn.  17  ;  stomach  of  Merluccius  merluccius, 
Stn.  8. 

This  parasite  has  been  recorded  from  numerous  fishes  in  polar  and  temperate 
waters  of  the  world.  This  appears,  however,  to  be  the  first  record  of  this  worm  from 
M.  macrophthalma,  T.  trachyrincus  and  M.  poutassou.  For  this  reason  a  table  of 
important  dimensions  is  given  (Table  3). 

ACKNOWLEDGMENTS 

My  thanks  are  due  to  the  following  :  Dr  J.  P.  Harding,  late  of  the  British  Museum 
(Natural  History),  and  Dr  A.  J.  Lee  of  the  M.A.F.F.  Fisheries  Laboratory,  Lowestoft, 
for  arranging  my  participation  in  the  cruise  ;  Mr  R.  J.  Blacker,  the  naturalist-in- 
charge,  and  other  M.A.F.F.  staff  and  crew  members  of  'Cirolana'  for  their  help  and 
advice  ;  Dr  N.  R.  Merrett  of  the  National  Institute  of  Oceanography,  Wormley, 
Surrey,  for  allowing  me  to  examine  fishes  in  the  N.I.O.  collection  ;  and  Mr  A. 
Wheeler  of  the  British  Museum  (Natural  History)  for  identifying  some  of  the  fish. 
Finally,  I  would  like  to  thank  Mr  S.  Prudhoe  for  his  advice  and  encouragement 
during  this  study. 

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1074  pp. 


R.  A.  BRAY 

Department  of  Zoology 

BRITISH  MUSEUM  (NATURAL  HISTORY) 

CROMWELL  ROAD 

LONDON  SW7  5BD 


A  LIST  OF  SUPPLEMENTS 
TO  THE  ZOOLOGICAL  SERIES 

OF  THE  BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 


1.  KAY,  E.  ALISON.     Marine  Molluscs  in  the  Cuming  Collection  British  Museum 
(Natural  History)  described  by  William  Harper  Pease.     Pp.  96  ;    14  Plates. 
1965.     (Out  of  Print.)     £3.75." 

2.  WHITEHEAD,  P.  J.  P.    The  Clupeoid  Fishes  described  by  Lacepede,  Cuvier  and 
Valenciennes.     Pp.  180  ;   n  Plates,  15  Text-figures.     1967.     £4. 

3.  TAYLOR,  J.  D.,  KENNEDY,  W.  J.  &  HALL,  A.  The  Shell  Structure  and  Mineralogy 
of  the  Bivalvia.      Introduction.      Nuculacea-Trigonacea.      Pp.  125  ;  29  Plates 
77  Text-figures.     1969.    £4.50. 

4.  HAYNES,  J.  R.     Cardigan  Bay  Recent  Foraminifera  (Cruises  of  the  R.V.  Antur) 
1962-1964.     Pp.  245  ;    33  Plates,  47  Text-figures.     1973.     £10.80. 


Printed  in  Great  Britain  by  John  Wright  and  Sons  Ltd.  at  The  Stonebridge  Press,  Bristol  BS4  sNU 


OBSERVATIONS  ON 
LINEARE  PENARD  (TESTACEA : 
PROTOZOA) 


R.  H.  HEDLEY 

AND 

C.  G.  OGDEN 


BULLETIN  OF 

THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 
ZOOLOGY  Vol.  26  No.  3 

LONDON:   1974 


OBSERVATIONS  ON  TRINEMA  LINEAR 
PENARD  (TESTACEA:  PROTOZOA) 


BY 

RONALD  HENDERSON  HEDLEY 

AND 

COLIN  GERALD  OGDEN  / 


Pp.  185-199  ;  7  Plates,  i  Text-figure 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
ZOOLOGY  Vol.  26  No.  3 

LONDON:  1974 


THE      BULLETIN     OF     THE      BRITISH     MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they 
become  ready.  Volumes  will  contain  about  three  or 
four  hundred  pages,  and  will  not  necessarily  be 
completed  within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
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Department. 

This  paper  is  Vol.  26,  No.  3  of  the  Zoological 
series.  The  abbreviated  titles  of  periodicals  cited 
follow  those  of  the  World  List  of  Scientific  Periodicals. 


World  List  abbreviation  : 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


Trustees  of  the  British  Museum  (Natural  History),  1974 


TRUSTEES  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

Issued  7  February,  1974  Price  £1.50 


OBSERVATIONS  ON  TRINEMA  LINEARE 
PENARD  (TE  STAGE  A  :  PROTOZOA) 

By  R.  H.  HEDLEY  AND  C.  G.  OGDEN 

CONTENTS 

Page 
SYNOPSIS  ...........          187 

INTRODUCTION  ...........         187 

Previous  work  -  biology  .          .          .          .          .          .          .          .188 

Previous  work  -  taxonomy       .          .          .          .          .          .          .          .188 

MATERIAL  AND  METHODS    .......          .          .          189 

DISTRIBUTION   ...........          190 

MORPHOLOGY  AND  VARIATION      ....  ...          191 

REPRODUCTION  ..........          192 

ULTRASTRUCTURE  OF  VEGETATIVE  STAGE       ...  .          192 

ACKNOWLEDGEMENT  ..........          197 

REFERENCES     ...........         197 

SYNOPSIS 

Trinema  lineave,  a  geographically  widespread,  fresh-water,  moss  and  soil  inhabiting  testacean, 
has  been  established  in  clonal  culture  with  a  doubling  time  of  between  72  and  78  hours.  Full 
descriptions  are  given  of  the  siliceous  shell  and  the  fine  structure  of  the  vegetative  stage.  Ultra- 
structural  features  of  special  interest  include  the  presence  of  microbodies,  and  a  microtubule- 
organizing-centre  associated  with  the  nucleus  at  prophase. 

INTRODUCTION 

THE  five  known  species  of  Trinema  Dujardin,  1841,  are  amongst  the  most  widely 
reported  rhizopods  to  be  found  in  soil,  sphagnum  moss  and  fresh-water  habitats. 
Very  little  is  known  of  their  biology  and  there  has  been  no  previous  report  of  the 
cytoplasmic  ultrastructure.  The  present  paper  is  an  account  of  the  fine  structure  of 
cultured  specimens  of  Trinema  lineare  Penard,  1890,  isolated  from  a  sample  of  moss 
and  soil  from  Cliffe  Marshes,  Rochester,  Kent,  in  February,  1970.  It  is  the  second 
of  a  series  of  papers  devoted  to  testate  amoebae  aimed  at  providing  a  fuller  under- 
standing of  their  biology,  and  a  fuller  appreciation  of  the  significance  and  nature  of 
the  siliceous  shell.  The  first  paper  in  the  series  was  devoted  to  a  detailed  account  of 
Euglypha  rotunda  -  a  widely  distributed  species  (Hedley  and  Ogden,  1973). 

Trinema  belongs  to  the  family  Euglyphidae  and  the  classification  adopted  here  is 
that  proposed  by  Loeblich  and  Tappan  (1961)  : 

Class  RHIZOPODEA        Von  Siebold,  1845 

Subclass  FILOSIA  Leidy,  1879 

Order  GROMIDA  Claparede  and  Lachmann,  1859 

Superfamily  EUGLYPHACEA  Loeblich  and  Tappan,  1961 


i88  R.    H.    HEDLEY    AND    C.    G.    OGDEN 

Family  EUGLYPHIDAE    Wallich,    1864; 

test  hyaline,  symmetrical,  elongate,  composed  of  rounded  siliceous 
scales,  aperture  rounded  or  elongate  :   one  nucleus. 

Previous  work  -  biology 

Leidy  (1879)  examined  several  testate  amoebae  from  North  America  arid  suggested 
that  the  cytoplasmic  structures  of  Trinema  were  similar  to  those  in  Euglypha,  in  that 
each  had  a  single  nucleus  and  two  contractile  vacuoles.  Penard  (1902)  observed 
that  when  the  animals  were  inactive,  or  in  the  vegetative  phase  of  their  life  cycle, 
the  granular  zone  and  nucleus  appeared  to  be  distinct.  Dunkerly  (1923)  reported 
that  reserve  shell-plates  were  arranged  around  the  nucleus,  and  suggested  that 
certain  dark  granules  were  chromidia.  Chardez  (1960)  noted  that  the  cytoplasm 
did  not  fill  the  shell  cavity  and  that  it  appeared  to  be  attached  only  at  the  apertural 
collar. 

Both  Penard  (1902)  and  Leidy  (1879)  described  the  pseudopodia  as  very  fine, 
usually  two  or  three  and  occasionally  six  in  number.  According  to  Leidy  (1879) 
when  the  animal  moves,  the  body  is  inclined  so  that  the  aperture  faces  anterior  and 
down,  while  the  fundus  points  backwards  and  up. 

During  binary  fission,  Penard  (1902)  observed  that  the  animals  are  diametrically 
opposed  whereas  at  conjugation  they  appear  to  be  directly  apposed.  Chardez 
(1960)  observed  the  part  of  the  reproductive  cycle  when  clear  cytoplasm  passed 
into  the  daughter  shell,  until  the  moment  that  the  daughter  cell  became  packed  with 
agitated  vacuoles.  The  cytoplasmic  volume  attained  full  size  within  20  minutes. 
Chardez  (1960)  also  reported  the  formation  of  a  cyst  with  two  nuclei  as  a  phenomenon 
of  conjugation,  in  which  the  cytoplasm  of  the  two  conjugants  become  joined  in  one 
shell,  the  empty  shell  remaining  provisionally  united.  Similar  united  individuals, 
with  one  shell  empty  and  the  other  containing  cytoplasm  and  two  nuclei,  were 
observed  by  Dunkerly  (1923)  who  considered  this  to  be  a  stage  of  encystation,  after 
which  the  cytoplasm  contracted,  the  chromidia  disappeared  -  at  least  as  staining 
bodies  -  and  finally  the  nuclei  fused.  Dunkerly  (1923)  also  suggested  that  the 
chromidia  were  used  up  during  encystation  as  reserve  food  material. 

As  a  result  of  an  ecological  study  of  a  Netherland  fen,  de  Graff  (1956)  reported 
that  Trinema  enchelys  and  T.  lineare  show  an  optimum  distribution  in  moderate 
dry  mosses,  but  were  found  in  most  kinds  of  biotopes,  only  T.  lineare  avoiding  the 
drier  mosses.  In  a  review  of  soil  protozoa,  Stout  and  Heal  (1967)  described  T.  lineare 
as  an  ubiquitous  species,  found  in  both  organic  and  mineral  topsoils,  and  having  the 
same  pH  tolerance  as  the  common  soil  ciliates  and  flagellates.  They  also  stated 
that  most  testaceans  -  with  some  exceptions  including  T.  lineare  -  reproduce  slowly, 
have  poor  encystment  mechanisms  and  poor  tolerance  to  high  carbon  dioxide  and 
low  oxygen  tensions  and  salinities. 

Previous  work  -  taxonomy 

T.  lineare  was  described  first  by  Penard  (1890),  who  considered  it  to  be  smaller 
and  more  elongate  than  T.  enchelys  (Ehrenberg,  1838).  The  same  author  (Penard, 


TRINEMA    LINEARE    (PROTOZOA)  189 

1902)  later  redefined  the  size  as  being  16-26  /urn  and  rarely  30  /un.  Cash  et  al.  (1915) 
redescribed  three  species  of  Trinema,  including  T.  lineare,  and  one  variety  in  a  review 
of  the  British  Rhizopoda  Fauna.  They  listed  T.  acinus  Leidy,  1879  as  a  synonym  of 
r.  lineare,  but  this  is  probably  a  doubtful  species  because  Leidy  (1879)  described  only 
T.  enchelys  and  figured  only  T.  acinus.  Both  Volz  (1929)  and  Kufferath  (1932) 
suggested  that  T.  lineare  was  a  synonym  of  T.  enchelys,  whereas  subsequent  authors 
have  recognized  both  species.  Hoogenraad  and  de  Groot  (1940)  redescribed  three 
species  and  listed  some  measurements  of  previous  authors.  Chardez  (1956)  figured 
numerous  variations  in  the  shape  and  size  of  the  shell  of  both  T.  enchelys  and  T. 
lineare. 

Thomas  (1958)  suggested  that  there  are  three  types  of  shell :  firstly,  those  with 
large  completely  overlapping  plates  ;  secondly,  those  with  large  incompletely  over- 
lapping plates  with  smaller  plates  filling  the  gaps  ;  and  thirdly,  those  with  large 
separate  plates  with  many  small  plates  ;  the  first  type  is  seen  fairly  often,  the  second 
is  very  common  whilst  the  last  type  is  rare. 

Bonnet  and  Thomas  (1960)  redescribed  five  species  of  Trinema  and  one  variety, 
T.  complanatum  Penard,  1890  ;  T.  complanatum  var.  globulosa  Chardez,  1959  ;  T. 
enchelys  ;  T.  galeata  (Penard,  1890)  ;  T.  lineare  and  T.  penardi  Thomas  and  Chardez, 
1958,  with  a  list  of  their  recorded  distribution.  In  several  publications  Decloitre 
(i96ia,  b,  I964a,  b,  I965a)  has  provided  additional  measurements  and  localities, 
and  discussed  variation  seen  in  some  testaceans.  Decloitre  (1962)  described  a  new 
variety  of  T.  lineare,  T.  lineare  var.  terricola  having  a  ventral  aperture,  and  more 
recently  Stepanek  (1967)  described  two  varieties,  T.  lineare  var.  globulosa  having  a 
mouth  at.  one  side  and  T.  lineare,  var.  pellucida  having  an  elliptical  mouth. 

MATERIALS  AND  METHODS 

T.  lineare  was  isolated  from  a  sample  of  moss  and  soil  collected  on  Cliffe  Marshes, 
near  Rochester,  Kent,  in  February,  1970.  Crude  cultures  were  made  from  small 
portions  of  this  material  placed  in  the  culture  liquid  and  kept  in  the  laboratory  at 
room  temperature,  i8-20°C.  Agnotobiotic  cultures  were  kept,  in  small  plastic 
containers,  on  a  thin  substrate  of  agar  (i  per  cent  agar  agar  in  distilled  water)  with  a 
sterilized  wheat  grain  added  prior  to  setting,  and  covered  with  a  shallow  layer  of  the 
culture  liquid.  This  liquid  was  a  5  per  cent  (w/v)  solution  of  soil  extract,  plus 
100  mg/l"1  of  sodium  nitrate  and  15  mg/1"1  of  sodium  dihydrogen  orthophosphate, 
in  distilled  water. 

Clonal  cultures  were  obtained  by  isolating  single,  active  animals.  One  such  clone 
was  subsequently  used  to  produce  working  cultures.  If  sub-cultures  are  made  at 
intervals  of  between  three  or  four  weeks  the  animals  appear  to  feed  and  reproduce 
readily.  The  clone  which  was  used  to  produce  working  cultures  is  now  deposited 
at  the  Culture  Centre  of  Algae  and  Protozoa,  the  Natural  Environment  Research 
Council,  Cambridge,  England. 

Optical  microscopy  -  The  animals  were  examined  by  bright-field  and  phase- 
contrast  microscopy,  either  alive  or  after  fixation.  Smears  fixed  in  either 
Schaudinn's  fluid  or  glutaraldehyde  were  stained  with  either  borax  carmine  or  iron 
haematoxylin. 


igo  R.    H.    MEDLEY    AND    C.    G.    OGDEN 

Scanning  electron  microscopy  -  For  morphological  studies  on  external  morphology 
both  live  animals  and  empty  shells  were  used.  Living  specimens  were  fixed  initially 
in  3  per  cent  glutaraldehyde  in  distilled  water  for  30  minutes.  The  empty  shells 
were  washed  initially  in  several  changes  of  distilled  water.  Single  specimens  were 
then  passed  through  several  changes  of  triple  glass  distilled  water  using  either  a 
single-hair  brush  or  a  fine-bore  pipette.  They  were  then  manipulated  onto  a  small 
cover-slip,  previously  cleaned  with  acetone  and  lint-free  tissue,  and  allowed  to  dry. 
Dried  specimens  adhere  well  to  glass  but  can  be  moved  by  use  of  a  moistened  single- 
hair  brush.  For  the  examination  of  individual  siliceous  plates,  single  clean  specimens 
were  placed  on  a  fragment  of  cleaned  cover-slip,  covered  with  a  small  drop  of  con- 
centrated sulphuric  acid  and  gently  heated.  On  evaporation  of  the  acid  the  plates 
are  liberated  from  the  organic  cement.  The  treated  cover-slips  were  attached  to 
'Stereoscan'  stubs  by  an  electrically-conductive  paint,  'Silver  Dag',  and  then  coated 
evenly  with  10-15  nm  °f  g°ld  using  an  Edwards  coating-unit  with  a  planetary 
specimen  holder  (Harris  et  aL,  1972).  The  stubs  were  examined  on  a  Cambridge 
Stereoscan  Mk  II  at  either  15  or  20  kV  and  the  results  recorded  on  Ilford  35  mm  H?3 
film. 

Transmission  electron  microscopy  -  Animals  were  fixed  at  room  temperature  for 
15  minutes  in  i  per  cent  glutaraldyhyde  in  0*05  M  Sorenson's  phosphate  buffer  plus 
0-0015  M  calcium  chloride,  followed  by  10  minutes  in  3  per  cent  glutaraldehyde  in  the 
same  buffer.  After  several  rinses  in  buffer,  they  were  post-fixed  in  i  per  cent  osmium 
textroxide  in  o-i  M  Sorenson's  buffer.  To  facilitate  subsequent  handling  the 
specimens  were  occasionally  embedded  in  1-5  per  cent  agar  at  this  stage,  prior  to 
dehydration.  The  material  was  dehydrated  by  passage  through  a  series  of  graded 
water/ethanol  mixtures,  ending  in  absolute  ethanol  and  embedded  in  Epon  812. 
Sections  were  cut  on  a  Porter  Blum  MT2  ultramicrotome  using  a  Du  Pont  diamond 
knife,  stained  with  alcoholic  uranyl  acetate  and  Reynold's  lead  citrate,  and  examined 
in  an  A.E.I.  6B  electron  microscope  operating  at  60  kV.  The  results  were  recorded 
on  Ilford's  'Special  Lantern  Contrasty'  plates. 


DISTRIBUTION 

T.  lineare  is  commonly  found  in  damp  and  wet  mosses  and  various  soils.  A  list  of 
localities  and  references  from  which  it  has  been  recorded  is  given  by  Bonnet  and 
Thomas  (1960).  Additional  localities  are  given  here  based  on  records  of  several 
authors,  namely,  Hoogenraad  and  de  Groot  (i952a,  b),  Bonnet  (1966),  Decloitre 
(i964a,  I965a,  b),  Chardez  (1961,  1969)  and  Golemansky  (1971). 

The  following  list  illustrates  the  geographically  wide  distribution  of  T.  lineare  : 


EUROPE  :  Great  Britain,  Spitzbergen,  Iceland,  Finland,  France,  Belgium, 

Netherlands,  Germany,  Italy,  Czechoslovakia  and  Hungary. 

NORTH  AMERICA  :  United  States  of  America,  Canada  and  Greenland. 
SOUTH  AMERICA  :   Tristan  da  Cunha,  Chile,  Bolivia  and  Peru. 


TRINEMA    LINEARE    (PROTOZOA)  191 

AFRICA  :  Angola,  Morocco,  Congo,  French  Cameroons,  Algiers,  Guinea, 

South  Africa  and  the  Seychelles. 

ASIA  :  India,  China,  Siberia  and  Japan. 

AUSTRALASIA  :        Australia,  New  Zealand  and  Tasmania. 
ANTARCTICA  :          South  Georgia  and  Gough  Island. 

MORPHOLOGY   AND   VARIATION 

The  siliceous  shell  is  elliptical  in  shape  through  the  major  axis  and  circular  through 
the  minor  axis  ;  it  varies  in  length  between  25  and  34-5  /mi  and  in  breadth  between 
14.5  and  19  /mi  (PI.  i,  figs.  B  and  E).  The  aperture  is  normally  circular,  invaginated 
and  situated  sub-terminally,  having  a  diameter  of  between  6  and  8-5  /mi  (PI.  i, 
figs.  A,  B  and  E).  Its  position  may  vary  greatly  through  various  angles  to  the 
longitudinal  axis  of  the  shell,  including  being  terminal.  The  shell  is  composed  of 
three  different  types  of  siliceous  plates  :  small,  circular  apertural  plates  (PI.  i, 
figs.  C  and  D),  large,  circular  shell-plates  (PI.  i,  fig.  B)  and  small,  oval  to  circular 
shell-plates  (PI.  i,  figs.  D  and  F). 

The  apertural  plates  are  roughly  circular,  folded  along  the  median  line  with  a  small 
central  dorsal  tooth  (PI.  i,  figs.  C  and  D),  and  vary  between  1-2  and  1-5  /mi  in  dia- 
meter. The  number  per  specimen  appears  to  vary  between  18  and  28,  but  the  majority 
of  animals  have  between  22  and  26.  The  apertural  plates  may  occasionally  be  moved 
from  their  marginal  position  either  to  form  double  rows  or  to  be  displaced  out  of 
symmetry.  Usually  either  one  or  two  rows  of  between  13  and  16,  small  shell-plates 
are  arranged  adjacent  to  the  apertural  plates  (PI.  i,  fig.  A),  but  these  are  also  subject 
to  some  variation,  in  one  instance  being  replaced  by  large  shell-plates. 

The  large  circular,  slightly  convex  shell-plates  have  a  diameter  of  between  4-0  and 
5-5  /mi  and  a  thickness  of  between  0-12  and  0-20  /mi ;  whereas  the  small  oval  to 
circular,  convex  shell-plates  (PI.  i,  fig.  F)  vary  in  length  between  2-4  and  3-5  /mi 
and  are  between  o-n  and  0-20  /mi  thick.  T.  lineare  has  approximately  50  incom- 
pletely overlapping  large  shell-plates  with  an  unknown  number  of  small  shell-plates 
haphazardly  filling  the  interstices,  many  of  them  being  completely  overlaid  by  the 
large  shell-plates.  This  is  the  second  and  most  common  of  the  three  types  of  shell- 
plate  arrangement  noted  by  Thomas  (1958).  Although  the  plates  are  normally 
arranged  evenly,  an  occasional  shell  is  seen  in  which  some  shell-plates  are  reversed, 
with  the  concave  surface  facing  outwards  (PI.  6,  fig.  E). 

The  incidence  of  abnormally  shaped  shells  (Plate  2)  is  usually  less  than  i  per  cent, 
although  as  many  as  10  per  cent  of  one  culture  was  observed  to  have  abnormal 
forms.  Cash  et  al.  (1915)  and  Chardez  (1970)  have  illustrated  previously  the  variation 
in  the  position  of  the  aperture  and  included  some  examples  of  evaginated  apertures. 
The  most  frequently  seen  abnormal  forms  are  a  single  shell  possessing  two  apertures  - 
an  aperture  being  defined  as  an  opening  bordered  by  apertural  plates.  Openings  in  the 
shell  due  to  inadequate  shell-plate  coverage  are  common  (PL  2,  fig.  F),  and  as  many  as 
six  openings  have  been  seen  in  one  individual.  It  would  appear  that  the  number  of 
shell-plates  in  abnormal  forms  is  roughly  proportional  to  the  number  of  apertures. 


IQ2  R.    H.    HEDLEY    AND    C.    G.    OGDEN 

There  is  also  a  slight  increase  in  the  amount  of  organic  cement  binding  the  plates  of 
the  abnormal  forms. 

There  appear  to  have  been  only  two  previous  reports  concerning  the  surface  ultra- 
structure  of  T.  lineare,  both  of  which  are  based  on  carbon  and  metal-shadowed  pre- 
parations. Thomas  and  Hovasse  (1962)  described  two  types  of  plates,  large  and 
small,  with  the  aperture  bordered  by  a  collar  of  biconvex  spines.  Mercier  et  al. 
(1964)  described  the  aperture  as  being  surrounded  by  one  or  two  circles  of  small  spiny 
plates. 

REPRODUCTION 

An  estimate  of  the  doubling  time  was  obtained  by  growing  three  replicate  cultures, 
and  recording  the  number  of  animals  present  at  regular  intervals.  Growth  curves 
produced  from  these  results  show  that  the  doubling  time  is  between  3-0  and  3-4  days. 
A  similar  calculation  for  Euglypha  rotunda  gave  a  doubling  time  of  between  1-4  and 
1-9  days  (Hedley  and  Ogden,  1973). 

Binary  fission  is  completed  in  approximately  60  minutes.  The  initial  stages  are 
difficult  to  observe  in  T.  lineare  due  to  the  small  size  of  the  animal  and  the  oblique 
position  of  the  aperture.  Cytoplasmic  division,  once  the  daughter  shell  has  been 
formed,  proceeds  in  a  manner  similar  to  that  previously  described  (Hedley  and 
Ogden,  1973)  for  E.  rotunda. 

ULTRASTRUCTURE   OF  VEGETATIVE   STAGE 

The  cytoplasm  does  not  quite  fill  the  shell  cavity  (PI.  3,  fig.  A),  but  appears  to  be 
anchored  to  the  apertural  collar.  Numerous  fine  processes  can  be  seen  extending 
between  the  cytoplasm  and  the  inner  shell  wall  (PI.  3,  fig.  A).  Unevenly  spaced 
pellicular  microtubules  lie  beneath  the  plasmalemma  and  run  in  an  antero-posterior 
direction.  The  mitochondria  are  ovoid  or  spherical  in  shape,  possess  tubular  cristae, 
and  appear  to  be  distributed  at  random  throughout  the  cytoplasm. 

Nucleus.  The  vegetative  or  interphase  nucleus  is  usually  spherical,  between 
4-4  and  5 -7  /nm  in  diameter,  occupying  a  central  position  at  the  posterior  end  of  the 
cytoplasm  (PI.  3,  figs.  A  and  D).  It  is  bound  by  a  nuclear  envelope  made  up  of  two 
tripartite  membranes,  the  outer  membrane  being  continuous  with  the  granular 
endoplasmic  reticulum.  The  nuclear  matrix  is  finely  granular,  with  small  con- 
centrations of  densely-staining  chromatin  scattered  throughout  and  a  dense  nucleolus. 

Although  the  nucleus  during  interphase  was  spherical  in  most  animals  examined, 
a  number  have  been  observed  which  are  cone-shaped  posteriorly  (PI.  4,  fig.  A),  and 
it  is  assumed  that  this  is  correlated  with  the  early  stages  of  prophase.  The  cone- 
shape  is  caused  by  the  convergence  of  numerous  microtubules  towards  a  specific 
region  at  the  posterior  end  of  the  cytoplasm.  The  microtubules  are  first  seen  con- 
verging on  the  nucleus  around  its  equatorial  region  (Text-fig,  i  ;  PI.  6,  fig.  A),  whilst 
at  a  level  in  the  region  of  the  contractile  vacuoles  they  lie  in  the  endoplasmic  reticulum 
at  some  distance  from  the  nucleus.  Although  it  is  difficult  to  estimate  the  number 
of  microtubules  present  because  of  the  close  relationship  of  the  dense  endoplasmic 
reticulum,  it  is  apparent  that  the  numbers  increase  to  approximately  70  in  the  pos- 
terior region  of  the  cytoplasm. 


TRINEMA  LINEARE  (PROTOZOA) 


193 


---B 


C- 


E-- 


MTOC 


(a) 


r--D 


(b) 


E--  - 


-F 


FIG.  i.  (a)  Diagram  of  a  longitudinal  section  through  the  nucleus  (see  PI.  4,  fig.  A), 
(b)  Tranverse  sections  at  those  levels  of  the  nucleus  marked  A-B,  C-D  and  E-F  in  (a), 
to  illustrate  the  microtubules  converging  towards  the  microtubule-organizing- centre 
(MTOC),  as  discussed  in  the  text. 


194  R-    H-    HEDLEY    AND    C.    G.    OGDEN 

The  microtubules  lie  close  to  the  nuclear  membrane  (PI.  4,  fig.  C),  but  do  not  lie 
in  invaginations  similar  to  those  reported  by  Leadbeater  and  Dodge  (1967)  for  the 
dinoflagellate  Woloszynskia  micra.  They  appear  to  attach  to  an  electron-dense  area 
close  to  the  distal  end  of  the  nucleus  (PI.  4,  figs.  C  and  D),  which  is  comparable  with 
previously  described  and  similar  regions  in  the  alga,  Char  a  (Pickett-Heaps,  1968), 
fungal  zoospores  (Fuller  and  Calhoun,  1968),  the  soil  amoeba,  Acanthamoeba  castel- 
lanii  (Bowers  and  Korn,  1968),  the  marine  amoebae,  Stereomyxa  ramosa  and  5. 
angulosa  (Benwitz  and  Grell,  igyia,  b),  and  the  marine  protist,  Labyrinthula  (Perkins, 
1970).  Such  regions  of  attachment  for  microtubules  are  referred  to  in  other  cells 
under  a  variety  of  terms.  For  example,  in  dividing  plant  cells,  Pickett-Heaps  (1969) 
referred  to  it  as  a  '  microtubule-organizing-centre '  -  MTOC,  and  suggested  that  such 
centres  initiate  and  control  the  arrangement  of  microtubules.  An  MTOC  was  also 
described  by  McCully  and  Robinow  (1972)  in  association  with  the  nucleus  during 
mitosis  in  yeasts.  In  Labyrinthula,  Perkins  (1970)  described  the  granular  aggregate 
as  a  'protocentriole',  while  the  term  '  paracentrosome '  was  suggested  by  Manton 
et  al.  (1970)  for  the  precursor  material  seen  to  accumulate  near  each  pole  at  meiotic 
division  in  the  marine  centric  diatom,  Lithodesium  undulatum.  Fuller  and  Calhoun 

(1968)  stated  that  the  kinetosome  of  fungal  zoospores  is  an  unlikely  'organizing 
centre'  but  suggested  that  the  electron-opaque  material  surrounding  the  proximal 
third  of  the  kinetosome  could  be  such  a  'centre'.     In  mouse  oocytes  electron-dense 
fibrillar  areas,  close  to  the  nucleus,  from  which  microtubules  radiate  are  referred  to 
by  Szollosi  et  al.  (1972)  as  'microtubule  foci'.     Evidence  that  such  regions  indicate 
the  site  of  microtubule  formation  in  many  systems  has  been  provided  by  Tilney  and 
Goddard  (1970)  and  Tilney  (1971),  who  conducted  experiments  on  the  breakdown  and 
reformation  of  microtubules  in  certain  species  of  Heliozoa. 

Contractile  vacuoles.  Two  or  three  contractile  vacuoles  occur  at  the  edge  of  the 
granular  endoplasmic  reticulum  (PI.  3,  fig.  D)  in  the  region  of  the  nucleus.  These 
vacuoles  are  often  surrounded  by  numerous  vesicles  which  are  associated  with  the 
lumen  of  the  vacuole,  and  they  discharge  directly  into  the  shell  cavity. 

Microbodies.  Microbodies  appear  in  all  the  specimens  examined  (PI.  3,  fig.  B). 
They  are  ovoid  or  spherical  in  shape,  varying  in  size  between  0-30  and  0-55  /mi, 
with  a  dense  granular  matrix  surrounded  by  a  single  unit  membrane.  Tubular 
elements,  between  18  and  26  nm  in  diameter,  appear  within  the  matrix  (PI.  4,  fig.  E), 
and  as  many  as  four  tubules  are  apparent  as  cross-sections  or  loops  in  some  micro- 
bodies. 

The  microbodies  in  T.  lineare  are  similar  to  those  reported  in  various  Foraminifera 
by  Hedley  et  al.  (1967),  Hedley  and  Wakefield  (1969)  and  Febvre-Chevalier  (1971), 
in  possessing  tubular-like  elements  within  the  matrix.  Hedley  and  Wakefield 

(1969)  imply  that  such  organelles  are  possibly  a  normal  component  of  the  cytoplasm 
of  the  Foraminifera.     The  structure  and  function  of  microbodies  has  been  reviewed 
recently  by  Hruban  and  Rechcigl  (1969). 

Endoplasmic  reticulum.  A  concentrated  mass  of  granular  endoplasmic  reticulum 
usually  surrounds  the  nucleus,  and  appears  more  electron-dense  than  the  surrounding 
cytoplasm  due  to  the  concentration  of  ribosomes  (PI.  3,  figs.  A  and  D).  In  early 


TRINEMA    LINEARE    (PROTOZOA)  195 

prophase,  however,  the  nucleus  is  drawn  out  distally  from  the  endoplasmic  reticulum 
region  (PI.  4,  fig.  A). 

Pigment  zone.  A  zone  of  large  vacuoles  lies  immediately  anterior  to  the  endoplas- 
mic reticulum  region  (PL  3,  fig.  A).  They  contain  electron-dense  material  and  have 
previously  been  equated  by  Hedley  and  Ogden  (1973)  to  the  'pigment  zone'  of  earlier 
light  microscopy  workers.  A  probable  developmental  sequence  in  the  formation  of 
the  electron-dense  material  in  the  vacuoles  is  illustrated  in  PI.  3,  figs.  A  and  B  -  the 
various  stages  are  labelled  i  to  4.  The  initial  stage  (i)  shows  the  matrix  to  be 
granular  with  a  small  electron-dense  area  in  the  centre.  As  this  deeply  stained  area 
increases  in  size  (2  and  3)  the  matrix  becomes  coarser  and  uneven  rents  appear  in  the 
electron-dense  material.  In  the  final  stage  (4)  the  vacuole  is  composed  mainly  of 
electron-dense  material.  The  rents  in  the  electron-dense  material  are  possibly 
caused  by  its  impermeability  to  the  embedding  resin.  No  explanation  can  be  made 
at  present  regarding  the  occurrence  of  this  constant  structure,  or  of  the  occasional 
empty  vacuoles  seen  here  and  reported  previously  by  Hedley  and  Ogden  (1973)  for 
Euglypha  rotunda. 

Golgi  apparatus.  A  single  Golgi  apparatus  lies  immediately  posterior  to  the  nucleus 
at  the  edge  of  the  endoplasmic  reticulum  region  (PI.  4,  fig.  A).  Both  smooth  and 
coated  vesicles  are  associated  with  the  Golgi  saccules  (PI.  4,  fig.  B).  In  addition,  the 
saccules  of  the  Golgi  are  often  distended  by  small  concentrations  of  densely  staining 
fibrillar  material  (PL  5,  fig.  C).  These  concentrations  appearing  in  the  outer  saccules 
of  the  dictyosome  become  progressively  spherical  as  the  concentration  of  material 
increases  and  finally  are  detached  at  the  margins  of  the  saccules  as  membrane- 
bound  vesicles.  At  this  stage  the  fibrillar  material  is  concentrated  mainly  at  the 
centre  of  the  vesicle,  and  has  small  strands  radiating  from  the  centre  (PL  3,  fig.  C). 
The  vesicles  then  pass  around  the  outside  of  the  endoplasmic  reticulum  region  and 
are  distributed  randomly  throughout  the  cytoplasm. 

It  has  already  been  established  (Favard,  1969)  that  the  Golgi  apparatus  appears 
to  play  a  role  in  the  packing  of  secretory  products  for  export  and  storage.  The 
chemical  constituents  of  such  secretory  products  in  both  plants  and  animals  are 
usually  polysaccharide  or  protein  macromolecules.  Hedley  and  Wakefield  (1969) 
suggested  that  the  polysaccharide  produced  by  the  Golgi  apparatus  of  the  marine 
protozoon,  Gromia  oviformis,  appeared  to  be  used  in  the  formation  of  the  proteinace- 
ous  shell-wall.  Schwab  (1969)  also  suggested  that  the  fibrillar  shell-wall  material 
in  the  marine  foraminifera,  Myxotheca  arenilega,  is  produced  by  the  Golgi  apparatus. 
More  recently,  Hedley  and  Ogden  (1973)  have  suggested  that  the  fine  fibrillar,  poly- 
saccharide material  contained  in  the  spherical,  membrane-bound  vesicles  of  E. 
rotunda  might  be  utilized  to  form  the  proteinaceous  cement  or  glue,  that  lines  the 
inside  of  the  siliceous  shell  and  also  binds  the  shell-plates  of  this  testate  amoeba. 

It  seems  reasonable  to  suggest  that  the  fibrillar  vesicles  (PL  3,  fig.  C)  produced  by 
the  Golgi  apparatus  in  T.  lineare  might  be  the  initial  stages  in  the  formation  of  the 
organic  cement  bodies  (PL  5,  fig.  E). 

Another  type  of  vesicle,  with  a  double  unit'  membrane  and  usually  electron- 
transparent  contents,  often  occurs  lying  in  the  area  of  cytoplasm  between  the  distal 


ig6  R.    H.    HEDLEY    AND    C.    G.    OGDEN 

end  of  the  granular  endoplasmic  reticulum  region  and  the  Golgi  apparatus  (PI.  5, 
fig.  D) .  Occasionally  they  occur  in  the  cytoplasm  anterior  to  the  granular  endoplas- 
mic reticulum  region,  surrounded  by  a  ring  of  smooth  endoplasmic  reticulum  (PI.  5, 
fig.  A).  The  function  of  these  vesicles  is  at  present  unknown. 

Food  particles.  Food  particles,  which  are  usually  gram-negative  bacteria,  occur 
throughout  the  cytoplasm,  whereas  in  E.  rotunda  they  are  confined  to  the  anterior 
third  of  the  cytoplasm  (Hedley  and  Ogderi,  1973).  Nevertheless,  digestion  in  T. 
lineare  probably  occurs  in  the  enlarged  food  vacuoles  which  are  found  normally  in 
the  anterior  cytoplasm. 

Reserve  shell-plates.  Reserve  shell-plates  are  formed  in  that  region  of  granular 
endoplasmic  reticulum  that  surrounds  the  nucleus  and  close  to  the  Golgi  apparatus. 
As  the  shell-plates  are  formed  they  become  closely  packed  together  in  a  region  just 
anterior  to  the  nucleus  (PI.  6,  fig.  D).  Both  large  and  small  shell-plates  appear  to  be 
formed  at  the  same  time.  The  apertural  plates  appear  to  be  the  last  to  be  formed 
and  are  usually  seen  only  in  the  posterior  region  of  the  cytoplasm,  with  their  dorsal 
teeth  pointing  outwards. 

Large  inclusions,  containing  electron-dense  material,  are  present  in  the  granular 
endoplasmic  reticulum  region  (PI.  3,  fig.  A).  These  inclusions  appear  to  be  less 
dense  than  the  '  pigment  zone '  vacuoles  and  are  often  seen  to  fuse  with  the  unit 
membrane  of  reserve  shell-plates  (PI.  6,  fig.  B).  It  seems  probable  that  these 
inclusions  are  associated  with  the  formation  of  the  siliceous  plates  as  they  are 
frequently  seen  in  specimens  containing  reserve  shell-plates.  It  is  noted,  however, 
that  whereas  in  sections  of  animals  fixed  and  subsequently  treated  in  the  absence  of 
heavy  metals  the  siliceous  plates  are  naturally  electron-dense  (PI.  7,  fig.  A),  the 
inclusions  in  the  endoplasmic  reticulum  are  electron-transparent. 

The  only  additional  reference  to  those  previously  reported  by  Hedley  and  Ogden 
(1973)  regarding  the  formation  of  siliceous  material  is  that  of  Cachon  and  Cachon 
(1971)  on  silica  metabolism  in  Radiolaria.  They  suggest  that  the  siliceous  shells  of 
digested  micro-organisms,  upon  which  radiolarians  feed,  are  used  to  produce  their 
siliceous  skeletons,  and  the  unused  siliceous  material  is  rejected  in  gel  form. 

Organic  cement.  The  organic  cement  or  glue  that  holds  the  plates  together  is  a 
fine  fibrillar  material  probably  produced  by  the  Golgi  apparatus.  It  is  circulated 
within  the  cytoplasm  in  membrane-bound  vesicles  (PI.  5,  fig.  E),  and  is  discharged 
mainly  in  the  anterior  region  where  the  vesicles  fuse  with  the  plasmalemma.  In 
abnormal  forms  there  is  usually  more  cement  at  shell-plate  junctions  than  in  normal 
specimens. 

Pseudopodia.  The  fine  structure  of  the  pseudopodia  consists  of  ground  plasm 
limited  by  a  membrane,  and  occasionally  containing  microfilaments.  Our  observa- 
tions are,  however,  limited  to  cytoplasmically  joined  individuals,  in  which  micro- 
tubules  are  often  seen  in  the  cytoplasm  internal  to  the  aperture,  but  not  externally 
(PI.  6,  fig.  C).  Similar  observations  on  microtubules  were  reported  for  Difflugiella 
sp.  by  Griffin  (1972)  and  for  E.  rotunda  by  Hedley  and  Ogden  (1973). 

Rosette  groups.  Rosette-like  groups  are  frequently  seen  in  clonal  cultures  which  are 
four  or  five  weeks  old  (PI.  2,  fig.  G  and  PI.  7,  figs.  C-E).  The  groups  are  usually 


TRINEMA    LINEARE    (PROTOZOA)  197 

composed  of  five  individuals  or  less  and  are  joined  by  cytoplasmic  connections. 
These  connections  contain  numerous  cement  bodies  and  less  frequently  mitochondria 
(PL  7,  fig.  C).  In  contrast  to  the  situation  reported  by  Hedley  and  Ogden  (1973), 
for  similar  formations  in  E.  rotunda,  microfilaments  are  seldom  present.  Individuals 
with  reserve  shell-plates  are  seldom  seen  in  rosette  groups,  suggesting  that  such 
formations  may  be  the  result  of  starvation. 

Abnormal  forms 

The  cytoplasm  of  animals  with  abnormal  shapes  (PI.  2)  is  similar  to  that  of  normal 
animals  with  the  exception  that  the  number  of  nuclei  appears  to  be  related  to  the 
number  of  apertures  (PI.  7,  fig.  F).  From  our  observations  all  attempts  by  these 
forms  to  divide  are  abortive. 

ACKNOWLEDGEMENT 

We  would  like  to  acknowledge  the  technical  assistance  of  Mrs  J.  E.  Dorahy. 

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rhizopoden  en  Heliozoen.     Leiden.     302  pp. 

I952(a).     Thekamobe  Moosrhizopoden  aus  Nordamerika.     Arch.  Hydrobiol.  47  :  229-262. 

I952(b).      Thekamobe   Moosrhizopoden   aus    Asien.      Arch.    Hydrobiol.   47   :    263-287. 

HRUBAN,    Z.    &    RECHCIGL,    M.     1969.     Microbodies   and   related   particles    :    morphology, 

biochemistry,    and   physiology.     Int.   Rev.    Cytol.   Suppl.    i.     Academic   Press,    London. 

296  pp. 

KUFFERATH,  H.     1932.     Rhizopodes  du  Congo.     Rev.  Zool.  Bot.  Afr.  23  :  52-60. 
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division  in  a  dinoflagellate.     Arch.  Mikrobiol.  57  :  239-254. 
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Survey  of  the  Territories.     Washington.     324  pp. 
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and  development  of  the  spindle  at  mitosis  and  meiosis  in  a  marine  centric  diatom  (Litho- 

desmium  undulatum).     IV.   The  second  meiotic  division  and   conclusion.     /.    Cell   Sci. 

7  : 407-443. 
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Rhodosporidium  sp.  (Rhodotorula  glutinis)  and  Aessosporon  salmonicolor  (Sporobolomyces 

salmonicolor}.     J.  Cell  Sci.  11:    1-31. 


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C.r.  Acad.  Sci.  (Paris).  258:  5967-5968. 
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Geneve,  31  :  1-230. 

1902.     Faune  Rhizopodique  du  Bassin  du  Ldman.     Geneva.     700  pp. 

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68  :  349-406. 


RONALD  HENDERSON  HEDLEY,  D.Sc. 
BRITISH  MUSEUM  (NATURAL  HISTORY) 
CROMWELL  ROAD 
LONDON  SW7  5BD 

COLIN  GERALD  OGDEN 

BRITISH  MUSEUM  (NATURAL  HISTORY) 

CROMWELL  ROAD 

LONDON  SW7  5BD 


PLATE    i 

A.  Aperture  of  T.  lineare  ;  note  the  invagination  produced  by  the  curved  shell-plates,    x  7100 

B.  Apertural  view  of  T.  lineare  showing  the  arrangement  of  large,  circular  shell-plates,    x  2900 

C.  Two  apertural  plates  ;   the  lower  one  shows  a  characteristic  median  fold.  x  23  ooo 

D.  A  preparation  from  which  the  organic  cement  has  been  removed  illustrating  the  arrange- 
ment of  apertural  and  small  shell-plates.  x  8650 

E.  Lateral  view  of  T.  lineare  illustrating  the  oblique  position  of  the  aperture.  x  2900 

F.  Individual  shell-plates  ;   note  the  fold  in  small  shell-plate  (arrowed).  x  n  500 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE    i 


PLATE   2 

A-F.  Abnormal  variants  of  T.  linear e  ;  note  that  the  apparent  imbrications  are  due  possibly 
to  electrons  penetrating  the  borders  of  the  shell-plates,  these  are  not  seen  when  the 
accelerating  voltage  on  the  scanning  electron  microscope  is  reduced. 

A,  B,  D,  E  and  F  x   1000     C   x  2000 

G.  Groups  of  three  animals  in  'rosette'  formation.  x   1400 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE 


PLATE   3 

A.  Longitudinal  section  showing  the  pigment  zone  (pz),  electron-dense  inclusion  (i),  granular 
endoplasmic  reticulum  (ger),  nucleus  (n)  and  Golgi  apparatus  (G).  x  4300 

B.  Section  showing  various  vacuoles  containing  bacteria  (b),  microbodies  (m),  and  the  probable 
stages  in  the  development  of  electron-dense  particles  (stages  1-3)  -stage  4  is  shown  in 
fig.  A.  x   15  950 

C.  Section  showing  vesicles  containing  fibrillar  material  (fm)  concentrated  in  the  centre  with 
radiating  strands.  x  23  900 

D.  Transverse  section  showing  three  contractile  vacuoles  (cv)  which  are  at  systole,  nucleus  (n) 
and  the  concentrated  mass  of  perinuclear  endoplasmic  reticulum  (ger).  x  7800 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE    3 


PLATE   4 

A.  Longitudinal  section  showing  the  position  and  ovoid  shape  of  the  nucleus  (n)  at  prophase, 
Golgi  apparatus  (G),  the  cytoplasmic  attachments  to  the  shell  and  the  concentrated  mass 
of  perinuclear  endoplasmic  reticulum  (ger.).  x  7800 

B.  Cross-section  of  the  posterior  region  of  nucleus  (n)  showing  proximity  of  numerous  micro- 
tubules  (arrowed),  Golgi  apparatus  (G)  and  coated  vesicles  (eves).  x  20  750 

C.  Section  showing  microtubule  (mt)  close  to  nuclear  membrane  (nm)  and  the  terminal  position 
of  the  microtubule-organizing-centre,  MTOC  (arrowed).  x  30  100 

D.  Section  immediately  posterior  to  the  nucleus  showing  nuclear  envelope  (ne)  and  numerous 
microtubules  (mt)  converging  onto  microtubule-organizing-centre,  MTOC  (arrowed). 

x  56  700 

E.  Section  through  two  microbodies  showing  tubular  elements.  x  44  600 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE 


PLATE   5 

A.  Vesicle  with  double-unit  membrane  and  enclosed  by  smooth  endoplasmic  reticulum  (ser) 
(see  also  fig.  D).  x  30  100 

B.  Transverse  section  through  anterior  region  of  cytoplasm  showing  numerous  cement  bodies 
(cm)  and  thin  cytoplasmic  strands  in  the  space  between  cytoplasm  and  shell.         x   10  400 

C.  Section  showing  Golgi  apparatus  (G)  with  concentrations  of  fibrillar  material  (fm)  in  the 
saccules,  nucleus  (n)  and  pellicular  microtubules  (pmt).  x  22  300 

D.  Probable  origin  of  electron-transparent  vesicles  in  the  region  between  granular  endoplasmic 
reticulum  (ger)  and  Golgi  apparatus  (G)  (see  also  fig.  A).  x  20  050 

E.  Cement  bodies  with  fibrillar  matrix  and  electron-dense  centres.  x  40  100 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE    5 


PLATE   6 

A.  Section    through    equatorial   region    of    nucleus    (n)    showing   surrounding   microtubules 
(arrowed).  x  30  100 

B.  Fusion  of  electron-dense  inclusion  with  reserve  shell-plate  vesicle.  x  22  300 

C.  Section  through  apertural  region  showing  internal  microtubules  (mt),  apertural  plate  (ap) 
and  pseudopodial  trunk  (pt)  which  is  relatively  structureless.  x   15  ooo 

D.  Stack  of  reserve  shell-plates  each  in  separate  membrane-bound  vesicle.  x  22  300 

E.  Portion  of  shell  showing  shell-plates  (arrowed)  with  concave  surface  outwards.          x  4300 


Bull.  Br.  Mm.  nat.  Hist.  (Zool.)  26,  3 


PLATE    6 


IP 


PLATE   7 

A.  Glutaraldehyde-fixed  and  unstained  section  showing  the  arrangement  of  reserve  shell-plates. 

x  4300 

B.  Section  of  two  animals  directly  apposed  ;    note  the  larger  individual  contains  numerous 
reserve  plates  and  a  well-defined  pigment  zone  (pz).  x  2100 

C.  Apertural  region  of  a  'rosette'  group  of  four  individuals  ;    note  the  presence  of  vacuoles 
containing  bacteria  (b)  and  the  numerous  cement  bodies  (cm).  x  4300 

D.  Section  through  two  united  adult  specimens  showing  cement  bodies  (cm)  and  pseudopodial 
extensions  ;   note  the  structureless  nature  of  the  pseudopodia.  x  7800 

E.  A  'rosette'  group  of  three  animals  showing  the  diffuse  nature  of  the  cytoplasm.        x  2800 

F.  Abnormal  individual  with  two  nuclei.  x  5750 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  3 


PLATE    7 


A  LIST  OF  SUPPLEMENTS 
TO  THE  ZOOLOGICAL  SERIES 

OF  THE  BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 


1.  KAY,  E.  ALISON.     Marine  Molluscs  in  the  Cuming  Collection  British  Museum 
(Natural  History)  described  by  William  Harper  Pease.     Pp.  96  ;    14  Plates. 
1965.     (Out  of  Print.)     £3-75." 

2.  WHITEHEAD,  P.  J.  P.     The  Clupeoid  Fishes  described  by  Lacepede,  Cuvier  and 
Valenciennes.     Pp.  180  ;   n  Plates,  15  Text-figures.     1967.     £4. 

3.  TAYLOR,  J.  D.,  KENNEDY,  W.  J.  &  HALL,  A.  The  Shell  Structure  and  Mineralogy 
of  the  Bivalvia.      Introduction.      Nuculacea-Trigonacea.      Pp.  125  ;  29  Plates 
77  Text-figures.     1969.     £4.50. 

4.  HAYNES,  J.  R.     Cardigan  Bay  Recent  Foraminifera  (Cruises  of  the  R.V.  Antur) 
1962-1964.     Pp.  245;   33  Plates,  47  Text-figures.     1973.    £10.80. 


Printed  in  Great  Britain  by  John  Wright  and  Sons  Ltd.  at  The  Stonebridge  Press,  Bristol  BS4  sNU 


22  JULJ 


THE  INDIGENOUS  EARTHWORMS 
(MEGASCOLECIDAE  :  OLIGOCHAETA) 

OF  TASMANIA 


B.  G.  M.  JAMIE  SON 


BULLETIN  OF 

THE    BRITISH    MUSEUM    (NATURAL    HISTORY) 
ZOOLOGY  Vol.  26  No.  4 

LONDON:   1974 


THE  INDIGENOUS  EARTHWORMS 

(MEGASCOLECIDAE  :  OLIGOCHAETA) 

OF  TASMANIA 


BY 

BARRIE  GILLEAN  MOLYNEUX  JAMIESON 

University  of  Queensland 


Pp.  201-328  ;   10  Plates  ;  32  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 

ZOOLOGY  Vol.  26  No.  4 

LONDON:  1974 


THE      BULLETIN      OF     THE     BRITISH      MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they 
become  ready.  Volumes  will  contain  about  three  or 
four  hundred  pages,  and  will  not  necessarily  be 
completed  within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  26,  No.  4  of  the  Zoological 
series.  The  abbreviated  titles  of  periodicals  cited 
follow  those  of  the  World  List  of  Scientific  Periodicals. 


World  List  abbreviation  : 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.). 


Trustees  of  the  British  Museum  (Natural  History),  1974 


TRUSTEES  OF 
THE  BRITISH  MUSEUM   (NATURAL  HISTORY) 

Issued  22  May,  1974  Price  £6.75 


THE  INDIGENOUS  EARTHWORMS 

(MEGASCOLECIDAE  :   OLIGOCHAETA) 

OF  TASMANIA 

By  B.  G.  M.  JAMIESON 

CONTENTS 

Page 

SYNOPSIS.         ...........  204 

INTRODUCTION  ...........  205 

EXPLANATION  OF  DESCRIPTIVE  FORMAT  AND  TERMINOLOGY      .          .          .  207 

SYSTEMATICS     ...........  212 

KEY  TO  THE  GENERA  OF  TASMANIAN  MEGASCOLECIDAE        .         .          .  212 
Subfamily  ACANTHODRILINAE  .          .          .          .          .          .          .          .212 

Tribe  Acanthodrilini 

Genus  Rhododrilus  Beddard  1889        .          .          .          .          .          .  212 

Rhododrilus  littoralis  sp.  n. .          .          .          .          .          .          .  213 

Subfamily  MEGASCOLECINAE    .         .         .          .          .         .         .         .  216 

Tribe  Perionychini       ......... 

Genus  Perionychella  Michaelsen,  1907  .          .          .          .          .  216 

Subgenus  Perionychella  Michaelsen,  1907  ....  220 

Subgenus  Vesiculodrilus  Jamieson,  1973  ....  221 

P.  (Perionychella)  capensis  sp.  n. .          .          .          .          .          .  225 

P.  (Perionychella)  hickmani  sp.  n.          .          .          .          .          .  226 

P.  (Perionychella)  irregularis  (Spencer,  1895)  •          •          •  22^ 

P.  (Perionychella)  kershawi  sp.  n.  .....  229 

P.  (Perionychella)  weldboroughi  sp.  n.    .          .          .          .          .  230 

P.  (Vesiculodrilus)  bithecata  sp.  n.          .          .          .          .          .  233 

P.  (Vesiculodrilus)  dilwynnia  (Spencer,  1895)          •          •          •  234 

P.  (Vesiculodrilus)  evansi  sp.  n.    .          .          .          .          .          .  236 

P.  ( Vesiculodrilus)  glandifera  sp.  n.       .....  237 

P.  (Vesiculodrilus)  hellyeri  sp.  n. .          .          .          .          .          .  238 

P.  (Vesiculodrilus)  hobartensis  (Spencer,  1895)         .          .          .  241 

P.  (Vesiculodrilus)  lacustris  (Stephenson,  1924)       .          .          .  245 

P.  (Vesiculodrilus)  montisarthuri  sp.  n.  ....  246 

P.  (Vesiculodrilus)  mortoni  (Spencer,  1895)    ....  247 

P.  ( Vesiculodrilus)  obliquae  sp.  n.  .          .          .          .          .  250 

P.  (Vesiculodrilus)  pedderensis  sp.  n.  .          .          .          .  251 

P.  (Vesiculodrilus)  tunnackensis  sp.  n.  .          .          .          .          .  253 

P.  (?)  bassana  (Spencer,  1895)      ......  253 

P.  (?)  decatheca  (Michaelsen,  1910)         .....  255 

P.  (?)  ellisi  (Spencer,  1895)  ......  257 

P.  (?)  moroea  (Spencer,  1895)       ......  257 

P.  (?)  richea  (Spencer,  1895)          ......  258 

P.  (?)  scolecoidea  (Spencer,  1895) ......  259 

Genus  Graliophilus  Jamieson,  1971     .          .          .          .          .          .  259 

Graliophilus  tripapillatus  sp.  n.    .          .          .          .          .          .  261 

Genus  Pinguidrilus  gen.  nov.      .......  263 

Pinguidrilus  tasmanianus  (Fletcher,  1888)     ....  263 

Genus  Woodwardiella  Stephenson,  1925        .....  265 

W oodivardiella  ?  tessellatus  (Spencer,  1895)     ....  266 


204  B.   G.   M.   JAMIESON 

Genus  Cryptodrilus  Fletcher,  1887       ......  266 

Cryptodrilus  albertisi  (Cognetti,  1910)  .....  270 

Cryptodrilus  avesiculatus  sp.  n.     .          .          .          .          .          .  271 

Cryptodrilus  brunyensis  sp.  n.       ......  273 

Cryptodrilus  dartnalli  sp.  n.  .          .          .          .          .          .  275 

Cryptodrilus  enter onephricus  sp.  n.         .....  277 

Cryptodrilus  ?  officeri  Spencer,  1895      .....  280 

Cryptodrilus  polynephricus  Spencer,  1895       ....  282 

C.  polynephricus  polynephricus  Spencer,  1895      .          .          .  284 

C.  polynephricus  urethrae  subsp.  n.    .          .          .          .          .  288 

C.  p.  polynephricus  ad  urethrae          .          .          .          .          .  291 

Cryptodrilus  simsoni  Spencer,  1895       .....  293 

Genus  Pseudocryptodrilus  Jamieson,  1972    .....  296 

Pseudocryptodrilus  acanthodriloides  sp.  n.  298 
Tribe  Dichogastrini 

Genus  Hickmaniella  gen.  nov.    .......  300 

Hickmaniella  opisthogaster  sp.  n.  .....  301 

Tribe  Megascolecini 

Genus  Oreoscolex  Jamieson,  1973         ......  302 

Oreoscolex  bidiverticulatus  sp.  n.   .          .          .          .          .          .  305 

Oreoscolex  campestris  (Spencer,  1895)    .          .          .          .          .  307 

Oreoscolex  irregularis  (Spencer,  1895)    .....  309 

Oreoscolex  leai  (Michaelsen,  1910)  .          .          .          .          .  310 

Oreoscolex  longus  sp.  n.        .          .          .          .          .          .          .  311 

Oreoscolex  peculiaris  sp.  n.  .          .          .          .          .          .          .  313 

Oreoscolex  sexthecatus  sp.  n.  .          .          .          .          .          .  315 

Oreoscolex  wellingtonensis  (Spencer,  1895)       .          .          .          .  317 

Genus  Megascolex  Templeton,  1844    .          .          .          .          .          .  318 

Megascolex  burniensis  sp.  n.          .          .          .          .          .          .  319 

Megascolex  montisarthuri  sp.  n.    .          .          .          .          .          .  321 

Megascolex  tasmanicus  (Spencer,  1895)  ....  324 

ACKNOWLEDGEMENTS          .          .          .          .          .          .          .          .          .  326 

REFERENCES     ...........  327 


SYNOPSIS 

This  paper  raises  the  number  of  indigenous  earthworms  known  from  Tasmania  to  forty-eight, 
in  ten  genera.  The  genus  Perionychella  Michaelsen,  i9O7a,  is  re-established  and  extended  to 
include  lumbricine  species  with  or  without  nephridial  bladders  and  perichaetine,  vesiculate 
species.  The  other  species  resemble  the  type-species  in  being  perichaetine  with  avesiculate 
nephridia.  All  vesiculate  species  are  included  in  the  subgenus  Vesiculodrilus  Jamieson,  I973a, 
initially  established  as  a  genus  for  lumbricine,  vesiculate  species.  Perionychella  absorbs  all 
Australian  species  of  Diporochaeta,  which  is  now  regarded  as  endemic  in  New  Zealand,  and  a 
large  portion  of  the  residue  of  Plutellus,  a  genus  which  has  been  revised  elsewhere.  A  check 
list  of  the  two  subgenera  of  Perionychella  is  given  which  lists  twenty-three  Tasmanian  species. 
Perionychella  is  therefore  the  dominant  genus  in  numbers  of  species  in  Tasmania,  as  in  Victoria. 
Twelve  of  the  Tasmanian  Perionychella  species  are  new  species  and  a  further  five  species  are  re- 
described,  three  of  these  from  new  material.  Only  P.  scolecoidea  (Spencer,  1895)  was  previously 
referred  to  the  genus  (Michaelsen,  i9O7a). 

The  remaining  earthworm  fauna  consists  of  a  new  species  and  the  first  Tasmanian  record  of 
Rhododrilus  ;  a  new  species,  also  the  first  record,  of  Graliophilus  ;  the  new  genus  Pinguidrilus 
erected  for  Notoscolex  tasmanianus  Fletcher  ;  a  species  of  Woodwardiella  which,  like  the  Gralio- 
philus species,  may  be  referable  to  Perionychella  ;  eight  species  of  Cryptodrilus,  of  which  four 


TASMANIAN  EARTHWORMS  205 

and  one  subspecies  are  new  ;  a  new  species  of  the  formerly  monotypic  Victorian  genus  Pseudo- 
cryptodrilus  ;  a  new  monotypic  genus  Hickmaniella  ;  eight  species  of  Oreoscolex,  of  which  four 
are  new,  and  three  species  provisionally  referred  to  Megascolex,  of  which  two  are  new.  There 
is  almost  total  specific  endemicity  in  Tasmania,  a  single  species,  Megascolex  tasmanicus,  being 
questionably  a  junior  synonym  of  a  mainland  species.  In  contrast  only  two  of  the  ten  genera 
are  endemic,  the  remaining  eight  occurring  on  the  mainland  and  five  of  these  in  Victoria,  the 
region  with  the  highest  zoogeographic  affinity  with  Tasmania.  Noteworthy  features  of  the 
Tasmanian  fauna  are  the  high  species  diversity  relative  to  the  area  of  the  island  (paralleled, 
however,  in  Victoria),  the  high  proportion  of  primitive,  chiefly  perionychin  genera  and  species  ; 
and  radiative  evolution  in  the  excretory  system  within  the  perionychin  genus  Cryptodrilus 
which  has  resulted  in  convergent  acquisition  of  grades  of  organization  characteristic  of  the  tribes 
Dichogastrini  and  Megascolecini,  these  tribes  being  poorly  represented  in  Tasmania. 


INTRODUCTION 

THE  earthworms  of  Tasmania  have  not  been  reviewed  since  the  original  work  of 
Spencer  (1895).  As  a  result  of  the  kind  cooperation  of  Mr  A.  J.  Dartnall  of  the 
Tasmanian  Museum  and  Art  Gallery,  Hobart,  and  of  Dr  J.  L.  Hickman  of  the 
Zoology  Department,  University  of  Tasmania,  in  placing  their  oligochaete  collec- 
tions at  the  author's  disposal,  the  taxonomic  and  zoogeographic  relationships  of  the 
Tasmanian  earthworm  fauna  have  been  investigated  and  the  number  of  known 
species  from  that  island  has  more  than  doubled.  The  present  paper  is  a  report  of 
these  investigations. 

Before  the  study  was  begun,  the  known  oligochaete  fauna  of  Tasmania  consisted 
of  six  species  of  aquatic  microdriles,  twenty-three  indigenous  species  of  megascolecid 
earthworms,  and  introduced  earthworms  of  the  family  Lumbricidae.  The  microdriles 
are  the  cosmopolitan  and  probably  anthropochorous  Lumbriculus  variegatus  (Muller, 
1774)  (Lumbriculidae)  ;  the  endemic  Telmatodrilus  multiprostatus  and  T.  pectinatus, 
both  of  Brinkhurst  1971  ;  Antipodrilus  davidis  (Benham,  1907)  which  also  occurs  in 
Australia  and  New  Zealand  ;  the  cosmopolitan  Limnodrilus  udekemianus  Claparede, 
1862  (all  Tubificidae)  ;  and  the  species  inquirenda,  Tasmaniaedrilus  tasmaniaensis 
Goddard  and  Malan,  1913  (Phreodrilidae) . 

The  previously  known  indigenous  earthworms  are  listed  below  under  their  original 
binomina.  Generic  identifications  were  rarely  correct  but  the  species  are  reassigned 
and  all  previous  generic  attributions  are  listed  in  the  specific  synonymies. 

Species  of  Spencer,  1895  : 

Cryptodrilus  irregularis  Megascolides  simsoni 
polynephricus  bassanus 

mortoni  Perichaeta  tasmanica 
hobartensis  moroea 

campestris  richea 

tesselatus  dilwynnia 

insularis  scolecoidea 

ellisi  irregularis 

wellingtonensis 
officeri 


144°         145° 


146° 


147° 


148° 


145° 


146 


147° 


148° 


FIG.  i.  Tasmanian  records  of  Perionychella  species.  0,  present;  Q.  absent;  ba,  P. 
bassana  ;  bi,  P.  (V.)  bithecata  ;  c.  P.  (P.)  capensis  ;  d,  P.  (V.)  dilwynnia  ;  el,  P.  ellisi  ; 
ev,  P.  (V.)  evansi  ;  g,  P.  (V.)  glandifera  ;  he,  P.  (V.)  hellyeri  ;  hi,  P.  (V.)  hickmani  ;  ho, 
P.  (V.)  hobartensis  ;  i,  P.  (P.)  irregularis  ;  k,  P.  (P.)  kershawi  ;  1,  P.  (P.)  lacustris  ;  mon, 
P.  (V.)  montisarthuri  ;  mor,  P.  moroea  ;  mort,  P.  (F.)  mortoni  ;  o,  P.  (F.)  obliquae  ;  p, 
P.  (F.)  pedderensis  ;  r,  P.  richea  ;  s,  P.  scolecoidea  ;  t,  P.  (F.)  tunnackensis  ;  w,  P.  (P.) 
weldboroughi.  Not  mapped  :  P.  decatheca.  (Numerals  pertain  to  map  references  in  text.) 


TASMANIAN  EARTHWORMS  207 

Further  species : 

Notoscolex  tasmanianus  Fletcher,  i888b         Notoscolex  leai  Michaelsen,  1910 
Megascolides  albertsi  Cognetti,  1910  Perionyx  lacustris  Stephenson,  1924 

Plutellus  decatheca  Michaelsen,  1910 


EXPLANATION  OF  DESCRIPTIVE  FORMAT  AND  TERMINOLOGY 

Those  unfamiliar  with  oligochaete  morphology  and  terminology  are  referred  to 
Stephenson  (1930)  for  what  remains  the  most  comprehensive  account  of  this  subject. 
It  should  be  noted,  however,  that  the  taxonomy  employed  by  Stephenson  is  now 
largely  obsolete.  In  the  following  explanation  of  the  format  used  in  describing 
species  in  the  present  work  some  additional  clarification  of  terminology  will  be  given. 

The  descriptive  format  has  been  kept  brief.  Some  characters,  e.g.  septal  thicken- 
ing, which  are  customarily  described  but  are  of  limited  value  are  omitted  in  favour  of 
fuller  treatment  of  systems  of  major  significance  in  taxonomy  which  are  frequently 
neglected,  such  as  the  excretory  system.  Some  information  retrievable  from  the 
illustrations,  e.g.  spermathecal  proportions  and  dimensions,  is  excluded  from  the 
text. 

These  preliminary  accounts  are  primarily  limited  to  a  holotype  and  paratype  but 
variation,  particularly  in  the  highly  species-specific  genital  fields,  is  appended.  For 
species  with  eight  setae  per  segment,  setal  ratios,  which  are  population-specific 
(Jamieson  and  Bradbury  1972),  are  given  for  several  specimens  where  these  are 
available.  Specimens  are  sexual,  and  usually  clitellate,  unless  otherwise  stated. 

Characters  employed,  in  sequence  are  as  follows. 

Dimensions  :  1  =  length,  w  =  midclitellar,  or  greatest  width  where  accounts  are 
drawn  only  from  the  literature,  s  =  number  of  segments.  These  and  other  numerical 
data  are  given  in  the  sequence  holotype  and  paratype  I.  Prostomium  whether 
epilobous  or  tanylobous  and  canaliculate  (with  middorsal  longitudinal  groove  or  not) . 
Body  form  is  mentioned  only  where  the  cross  section  departs  from  an  approximate 
circle.  The  anus  is  terminal  in  all  cases.  Position  of  the  first  dorsal  pore  :  deter- 
mined by  parting  the  intersegmental  furrows  with  a  probe  or  observing  expulsion  of 
alcohol  when  the  specimen  is  immersed  temporarily  in  water. 

Setal  ratios  (intersetal  distances)  :  the  four  setae  on  each  side  in  a  segment  are 
designated,  in  the  ventral  to  dorsal  direction,  a,  b,  c  and  d.  The  distances  aa  :  ab  : 
be  :  cd  :  dd  :  dc  :  cb  :  ba,  measured  by  camera  lucida,  are  expressed  in  the  tables  rela- 
tive to  a  constant  total  (periphery,  u)  of  100.  Direct  comparison  of  ratios  between 
individuals  without  recourse  to  additional  computations  is  also  facilitated  by  stating 
the  distances  as  ratios  relative  to  one  of  the  distances  (ab)  expressed  as  unity.  Where 
there  are  more  than  eight  setae  per  segment  (the  perichaetine  condition),  the  longi- 
tudinal setal  rows  are  designated,  in  the  ventral  to  dorsal  direction,  setal  line  I,  2, 
3,  etc.  but  the  ventral  two  rows  are  alternatively  termed  a  and  b.  The  dorsalmost 
row,  so  as  readily  to  indicate  this  position,  is  termed  setal  line  z  and  the  penultimate 
and  next  ventral  rows  are  y  and  x  respectively.  Setae  commence  on  segment  II 
in  all  described  species. 


208 


B.   G.   M.   JAMIESON 


144°          145" 


146° 


147° 


148° 


40 


O  polynephricus  polynephricus 
O  polynephricus  urethrae 

p. polynephricus  ad  urethrae 

enteronephricus 


avesiculatus, 
dartnalli 


145° 


146° 


147° 


148° 


FIG.  2.     Tasmanian  records  of  Cryptodrilus  species  and  subspecies.     0,  present ; 
Oi  absent.  (Numerals  pertain  to  map  references  in  text.) 


TASMANIAN   EARTHWORMS  209 

Nephropores  in  holonephric  Tasmanian  species  are  invariably  near  the  anterior 
margins  of  their  segments.  Their  position  relative  to  the  setal  lines  is  recorded. 

The  clitellum  may  be  annular,  i.e.  completely  encircling  the  body,  or  saddle-shaped, 
i.e.  interrupted  ventrally  throughout  its  extent.  Segmentation  indicated  as,  for 
instance,  XIII-XVII,  indicates  that  the  clitellum  extends  throughout  the  mentioned 
segments,  in  this  case  from  the  anterior  border  of  the  thirteenth  to  the  posterior 
border  of  the  seventeenth  segment.  A  fractional  indication,  such  as  ^XIII-^XVII, 
indicates  the  position  on  the  segment  at  which  the  clitellum  begins  or  ends,  in  this 
case  the  clitellum  would  include  the  posterior  two-thirds  of  XIII  and  the  anterior 
third  of  XVII.  The  male  genital  field  is  described  next,  including  the  position  of  the 
porophores  bearing  the  combined  pores  of  the  vasa  deferentia  and  the  pair  of  prostate 
glands,  and  the  form  and  configuration  of  accessory  genital  markings  in  their  vicinity. 
These  markings  are  usually  protuberant,  and  where  an  elliptical  marking  has  a  dis- 
tinct central  area  are  described  as  eye-like.  Other  accessory  genital  markings  may 
be  present  near  the  spermathecal  pores.  Female  pores  are  without  exception  paired 
and  anteromedian  to  setae  a  of  segment  XIV  in  Tasmanian  species  of  Perionychella 
but  their  position  is  recorded  for  each  species  pending  description  of  at  least  the 
entire  Australian  fauna.  Spermathecal  pores  are  intersegmental  or  segmental  and 
paired  or  single  in  Tasmanian  species  ;  their  number,  intersegments  or  segments 
occupied  and  location  relative  to  setal  lines  are  indicated. 

The  internal  anatomy  (examined  after  middorsal  longitudinal  incision)  is  described 
in  the  following  sequence.  Blood  vascular  system  :  in  all  Tasmanian  species  the 
dorsal  vessel  is  continuous  onto  the  pharynx  and  a  subneural  vessel  is  absent ;  the 
segment  occupied  by  the  last  hearts  is  given  and  the  statement  that  they  are  latero- 
oesophageal  refers  to  their  origin  from  the  supra-oesophageal  vessel  (or  the  oesophageal 
plexus) .  A  latero-oesophageal  heart  usually  also  receives  a  connective  from  the  dorsal 
vessel  but  this  was  not  always  demonstrable  in  the  material  examined.  The  extent 
of  the  supra-oesophageal  vessel  is  indicated  where  precisely  determinable  ;  the  stated 
degree  of  development  of  the  vessel  refers  to  the  maximum  development  in  its  course. 
The  account  of  the  alimentary  canal  gives  the  segmental  location  and  degree  of 
development  of  the  gizzard  ;  modification  of  the  oesophagus,  including,  where 
feasible,  dilatation  and  internal  specializations  such  as  rugae  and  lamellae,  and  the 
position,  if  present,  of  extramural  oesophageal  (calciferous  ?)  'glands'  ;  the  segment 
of  origin  of  the  intestine  (determined  by  intestinal  expansion  and,  where  possible, 
identification  of  the  oesophageal  valve  between  oesophagus  and  intestine)  and 
presence  or  absence  of  a  typhlosole.  Special  muscular  thickening  of  the  intestine 
is  present  only  in  Hickmaniella.  Intestinal  caeca  are  absent  in  all  Tasmanian 
species  of  the  Megascolecidae. 

The  types  of  nephridia  found  in  oligochaetes  are  described  by  Jamieson  (igyia). 
Nephridia  are  holonephridia  in  all  species  of  Perionychella,  Graliophilus,  Pinguidrilus, 
Woodwardiella  and  Rhododrilus  but  in  some  of  these  holonephric  species  anterior 
nephridia  are  tufted,  having  numerous  tubules,  with  simple  or  less  commonly  com- 
posite or  multiple  ducts  ;  it  remains  to  be  ascertained  whether  the  tufted  nephridia 
always  retain  the  preseptal  funnel  (the  stomate  condition)  of  the  simple  nephridia. 
The  most  anterior  segment  in  which  the  nephridia  were  seen  to  have  preseptal  funnels 


144°         145° 


B.   G.  M.   JAMIESON 
146°  147° 


148° 


145' 


146 


147° 


148° 


FIG.  3.     Tasmanian  records  of  Oreoscolex  species.     0,  present ;  Q.  absent.     Not  mapped  : 
Oreoscolex  leai.     (Numerals  pertain  to  map  references  in  text.) 


TASMANIAN  EARTHWORMS  211 

is  stated  but  it  is  not  to  be  assumed  that  funnels  were  absent  from  more  anterior 
nephridia  as  demonstration  of  funnels  is  difficult,  particularly  where  disruption  of 
other  organ  systems  is  undesirable.  Presence  or  absence  of  ectal  bladders,  the 
vesiculate  or  avesiculate  condition,  and  their  form  is  recorded,  as  is  the  occurrence 
of  diverticula  on  the  bladders. 

The  anterior  male  organs  are  described  next ;  whether  holandric  (testes  and  funnels 
paired  in  each  of  X  and  XI)  or  metandric  (these  organs  in  XI  only)  and  iridescence  of 
the  funnels  is  noted  (an  indication,  albeit  only  partly  reliable,  that  the  organs  are 
functional).  Location  of  seminal  vesicles  is  noted. 

Female  organs  in  Tasmanian  megascolecids  are  metagynous  (ovaries  and  funnels 
in  XIII)  as  in  all  but  a  few  species  of  the  suborder  Lumbricina  of  Brinkhurst  and 
Jamieson  1971.  This  and  the  presence  or  absence  of  ovisacs  (dependent  from  the 
anterior  wall  of  XIV)  is  noted  for  each  species. 

The  form  and  extent  of  the  prostate  glands  are  recorded  ;  variation  in  the  form  of 
these  is  discussed  in  the  introductory  discussion  of  Perionychella.  Penial  setae  : 
occurrence  of  these  enlarged  copulatory  setae  in  segment  XVIII,  associated  with  the 
male  pores,  is  noted.  Lengths  of  well-developed  setae  are  given  and  their  mor- 
phology as  revealed  by  the  light  microscope  and  scanning  electron  microscope  is 
briefly  described.  The  penial  setae  in  Tasmanian  species  of  Perionychella  are  un- 
usually slender  and  are  at  most  only  weakly  ornamented,  a  contrast  with  other 
genera  on  the  island,  and  reproduction  of  the  electron  microscope  photographs  is 
rarely  merited.  In  several  species  of  Perionychella  complete  or  undamaged  setae 
were  not  recoverable  and  the  morphology  could  not  be  elucidated.  A  study  of  the 
degree  of  inter-specific  variation  in  their  ultra-morphology  is  required. 

Finally,  the  number  of  spermathecae  and  the  form  and  number  of  their  diverticula 
are  noted,  detailed  information  being  retrievable  from  the  illustrations. 

Material  examined  :  Under  this  heading  are  given,  in  sequence,  the  locality  with 
habitat  data,  the  longitude  and  latitude  of  the  locality,  the  grid  reference  on  the 
map,  the  collector's  name,  the  date  of  collection,  and  the  institutions  in  which  the 
holotype,  paratypes  and  additional  specimens  are  lodged.  Letters  and  numbers 
following  the  abbreviation  for  the  institution  are  the  registration  numbers  of  the 
institution.  Lectotypes  and  paralectotypes  are  designations  of  Jensz  and  Smith 
(1969).  Abbreviations  for  the  institutions  are  given  below  and  listed  with  them 
are  the  names  of  collectors. 

Collectors  Institutions  in  which  specimens 

are  lodged 

Mr  E.  A.  Bradbury  AM  Australian  Museum,  Sydney 

Mr  A.  Dartnall  BJ  Author's  collections 

Dr  J.  W.  Evans  BM(NH)     British  Museum  (Natural 

Dr  J.  L.  Hickman  History) 

Professor  V.  V.  Hickman  NMV          National  Museum  of  Victoria 

Mr  W.  Jackson  TM  Tasmanian  Museum, 

Dr  B.  G.  Jamieson  Hobart 

Mr  R.  W.  Kerr 


B.   G.   M.   JAMIESON 


Collectors  (cont.) 

Mr  R.  C.  Kershaw 

Mr  C.  D.  King 

Mr  W.  Radford 

Tasmanian  Biological  Survey 

Mr  D.  Tyler 


SYSTEMATICS 


The  generic  and  specific  descriptions  are  arranged  under  the  relevant  subfamilies 
and  tribes  but  a  key  to  all  Tasmanian  genera,  ignoring  suprageneric  ranks,  will  be 
given  first.  This  key  is  intended  only  for  known  Tasmanian  species  of  the  genera 
concerned  and  should  not  be  used  as  a  general  diagnostic  key  for  these  genera  in 
other  regions. 

KEY  TO  THE  GENERA  OF  TASMANIAN  MEGASCOLECIDAE 

1  Combined  male  and  prostatic  pores  a  pair  on  XVII   (i6th  setigerous  segment). 

(Holonephric ;  bladders  absent)        ...  .          RHODODRILUS 

-  Combined  male  and  prostatic  pores  a  pair  on  XVIII  (lyth  setigerous  segment). 

(Holonephric  or  meronephric  ;   bladders  present  or  absent)  ....  2 

2  Nephridia  one  pair  per  segment  (holonephridia)  in  midbody   .....  3 

-  Nephridia  more  than  one  pair  per  segment  (meronephridia)  in  midbody  ...  6 

3  Nephridial  bladders  present         ..........  4 

-  Nephridial  bladders  absent  ..........  5 

4  Nephridia  discharging  in  irregular  rows,  from  c  to  near  middorsum.     Spermathecae 

adiverticulate PINGUIDRILUS 

-  Nephridia  discharging  in  a  straight  line,  in  the  vicinity  of  c  lines.     Spermathecae 

diverticulate .          ....  PERIONYCHELLA  (VESICULODRILUS) 

5  Intestinal  typhlosole  present       GRALIOPHILUS  (part ;   see  also  WOODWARDIELLA) 

-  Intestinal  typhlosole  absent          .          .          .    PERIONYCHELLA  (PERIONYCHELLA) 

6  Nephridia  2  or  3  on  each  side  except  at  the  caudal  extremity  which  is  holonephric 

PSEUDOCR  YPTODRILUS 

-  Nephridia  more  than  2  per  segment  throughout    .......  7 

7  Setae  more  than  8  per  segment   .......... 

Setae  8  per  segment  .........•••  9 

8  A   gizzard  present  at  the  beginning  of  the   intestine.     Oesophageal   gizzard   also 

present HICKMANIELLA 

-  Oesophageal  gizzard  present.     Intestinal  gizzard  absent          .          .  MEGASCOLEX 

9  Caudal  nephridia  with  numerous  funnels  on  each  side  on  the  anterior  face  of  each 

septum.     Bladders  absent        .......  OREOSCOLEX 

-  Caudal  nephridia  with  at  most  a  single  funnel  on  each  side  on  the  anterior  face  of  each 

septum.     Bladders  often  present CRYPT ODRILUS 

Subfamily  ACANTHODRILINAE 

Tribe  ACANTHODRILINI  s.  Jamieson,  igjiz 

Genus  RHODODRILUS  Beddard,  1889 

DIAGNOSIS.  Setae  8  per  segment.  Prostatic  pores  i  pair,  on  XVII  (in  one  species 
on  XVI)  ;  male  pores  combined  with  the  prostatic  pores  or  i  pair,  usually  on  XVII 


TASMANIAN   EARTHWORMS  213 

close  to  them,  sometimes  anterior  on  XVIII,  occasionally  associated  with  the  pos- 
terior ends  of  seminal  grooves.  Female  pores  i  pair,  on  XIV.  Spermathecal  pores 
1-5  pairs,  the  last  pair  anterior  to  9/10.  Nephropores  in  a  single  series  on  each 
side.  Last  hearts  in  XII  or  XIII.  Gizzard  single,  occupying  i  or  2  segments  in 
the  region  of  V-VIII,  usually  strongly  developed,  occasionally  small.  Holonephric 
throughout  or  exceptionally  with  anterior  meronephridia.  Nephridial  vesicles  of 
various  forms  or  absent.  Calciferous  glands  present  or  absent.  Holandric ; 
gymnorchous.  Prostates  tubular ;  penial  setae  usually  present.  Spermathecae 
diverticulate. 

TYPE-SPECIES.     Rhododrilus  minutus  Beddard,  1889. 

DISTRIBUTION.  New  Zealand  and  neighbouring  islands.  Two  species  in  Austra- 
lia :  one  in  Queensland,  the  other  in  Tasmania. 

TASMANIAN  SPECIES.    Rhododrilus  littoralis  sp.  n. 

REMARKS.  The  above  generic  definition  is  derived  from  that  of  Jamieson  (igjib). 
In  the  latter  work  attention  was  drawn  to  the  unsatisfactory  distinction  between 
Rhododrilus  and  Microscolex.  Although  the  majority  of  species  of  Microscolex  differ 
from  Rhododrilus  in  having  two  pairs  of  prostates,  the  type-species,  M.  phosphoreus 
(Duges),  like  M.  dubius  (Fletcher),  has  a  single  pair  of  prostates,  as  in  Rhododrilus, 
and  has  been  shown  by  Lee  (1970)  to  have  strong  phenetic  affinities  with  the  latter 
genus  (including  the  type-species,  R.  minutus)  over  which  it  has  chronological 
priority.  It  is  possible  that  Rhododrilus  will  have  to  be  synonymized  with  Micro- 
scolex and  new  genera  erected  for  some  of  the  species  of  the  present  Microscolex- 
Rhododrilus  complex  which  fall  outside  a  redefined  Microscolex.  The  following  new 
Tasmanian  species  is  provisionally  placed  in  Rhododrilus  because,  like  some  other 
species  of  this  genus,  including  the  type-species,  it  lacks  nephridial  bladders  whereas 
these  are  invariably  present  in  Microscolex.  In  location  of  nephropores  shortly 
below  c  lines  it  resembles  Microscolex  dubius,  however,  and  differs  from  Rhododrilus 
in  which  nephropores  are  in  b  or  c. 

Rhododrilus  littoralis  sp.  n. 
Fig.  4,  26C,  32E  ;  Plate  60.     Table  i 
1  =  65,  74  mm,  w  =  2  mm,  s  =  103,  105. 

Setal  annuli  strongly  protuberant,  excepting  the  anterior  preclitellar  and  extreme 
caudal  regions,  and  demarcated  posteriorly  by  a  groove  equalling  the  intersegmental 
furrows  in  distinctness.  Prostomium  epilobous  f,  \,  open,  margins  posteriorly 
convergent  but  indistinct.  Dorsal  pores  weakly  developed  behind  the  clitellum, 
possibly  present  as  rudiments  in  the  immediate  preclitellar  segments.  Setae  8  per 
segment,  commencing  on  II,  in  regular  longitudinal  lines  throughout ;  setae  a  and  b 
absent  (replaced  by  penial  setae)  in  XVII.  Nephropores  not  externally  visible. 
Clitellum  externally  indistinct,  annular,  £XIII-f  XVIII  (=  5|  segments)  from 
dorsal  incision  (holotype)  ;  weakly  developed  but  limits  indeterminable  in  paratype  i. 
Male  pores  in  XVII,  in  b  on  small  whitish  papillae  which  fill  ab  transversely  and 


2I4 


144°          145° 


B.  G.   M.   JAMIESON 
146°  147° 


148° 


145° 


146° 


147° 


148° 


FIG.  4.  Records  of  Tasmanian  megascolecid  earthworms  excluding  Perionychella,  Crypto- 
drilus  and  Oreoscolex  species.  •,  present ;  Q,  absent.  (Numerals  pertain  to  map 
references  in  text.) 


TASMANIAN  EARTHWORMS  215 

occupy  the  posterior  two-thirds  of  the  segment.  Accessory  genital  marking  a 
transverse  pad  in  19/20,  extending  laterally  shortly  beyond  b  lines,  and  longitudinally 
abutting  on  the  setal  annuli  of  XIX  and  XX  ;  strongly  tumid  and  transversely 
bisected  by  a  furrow  corresponding  with  but  not  peripherally  continuous  with  inter- 
segmental  furrow  19/20  (holotype)  ;  genital  marking  not  certainly  detectable  in 
paratype  I.  Female  pores  a  pair  on  XIV,  midway  between  the  setal  arc  and  an- 
terior margin,  very  slightly  lateral  of  a  lines  (paratype  i)  ;  not  certainly  demonstrable 
but  apparently  concealed  in  the  anterior  furrow  delimiting  the  setal  annulus  in 
holotype.  Spermathecal  pores  minute,  concealed  (holotype)  or  visible  with  difficulty 
(paratype  i)  in  intersegments  7/8  and  8/9,  in  b  lines. 

TABLE  i 

Intersetal  distances  in  segment  XII  in  Rhododrilus  littoralis 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

holotype  0-7  0-4  0-9  0-8  1-8  0-9  0-8  0-4 

paratype  I  0-8  0-5  0-9  0-8  1-8  0-8  0-9  0-5 

standardized  (u  —  100) 

aa  ab  be 

holotype 
paratype  i 
mean 
interval/aft 

Last  hearts  in  XIII  (those  in  X-XIII  latero-oesophageal ;  with  connectives  to 
the  dorsal  and  supra-oesophageal  vessel) .  Supra-oesophageal  vessel  in  VIII  ?  -XIV  ; 
moderately  developed.  Gizzard  small  and  easily  compressible,  though  muscular, 
in  V  (holotype)  ;  rudimentary  in  paratype  i.  Oesophagus  lacking  calciferous 
glands  ;  especially  vascularized  and  with  low  internal  rugae  in  VIII-XVI ;  slender 
and  less  vascular  in  XVII  and  XVIII ;  intestine  commencing,  with  dilatation,  in 
XIX  ?  (holotype,  paratype  i)  ;  the  alimentary  canal  in  XVII  and  XVIII  possibly 
intestinal,  however ;  typhlosole  absent.  Nephridia  st ornate  holonephridia,  com- 
mencing in  II,  preseptal  funnel  (demonstrated  in  intestinal  region)  large,  in  b  line  ; 
avesiculate  duct  entering  parietes  one-third  be  below  c.  Holandric  ;  gymnorchous  ; 
testes  large,  much  branched ;  sperm  funnels  small,  faintly  iridescent ;  seminal 
vesicles  racemose,  in  IX,  XI  and  XII  increasing  in  size  posteriorly.  Metagynous 
(ovaries  with  many  egg-strings)  ;  ovisacs  present  (paratype  i)  ;  not  detectable  in 
holotype.  Prostates  one  pair,  very  slenderly  tubular,  in  XVII-XXIII  (holotype) - 
XXIV  (paratype  i)  ;  vas  deferens  joining  the  muscular  duct  well  ectal  of  the  gland. 
Penisetal  follicles  conspicuous,  as  wide  as  the  prostates  and  almost  as  extensive, 
extending  from  XVII-XXII.  Penial  setae  filiform,  sinuous  ;  the  ectal  extremity 
depressed  spatulate  ;  ornamentation  absent ;  length  =  2-8-3-7  mm,  general  width 
of  shaft  =  ii  /mi  (paratype  i).  Spermathecae  3  pairs,  in  VII,  VIII  and  IX,  each 
with  ovoid  ampulla  and  shorter  digitiform  (uninseminated)  diverticulum  on  the 


aa 

ab 

be 

cd 

dd 

dc 

Cb 

ba 

10-00 

5'79 

13-16 

12-63 

27-36 

13-16 

12-11 

5'79 

11-22 

6-63 

13-27 

n-73 

26-02 

11-22 

12-76 

7-14 

10-61 

6-21 

13-22 

12-18 

26-69 

12-19 

12-44 

6-47 

1-71 

I-OO 

2-13 

1-96 

4-30 

I-96 

2-00 

1-04 

216  B.    G.   M.    JAMIESON 

short,  poorly  demarcated  duct ;    the  spermatheca  and  diverticulum  almost  sessile 
on  the  body  wall. 

Genital  markings.  Of  13  paratypes,  n  have  the  genital  field  described  for  the 
holotype,  with  a  single  accessory  genital  marking,  sometimes  only  faintly  indicated, 
at  19/20  ;  2  lack  the  marking  but  are  less  mature  than  the  others. 

MATERIAL  EXAMINED. 

Eagle  Hawk  Neck,  in  kelp  on  rocks,  I47°55'E.  43°oo'S.  Map.  19/2,  Dr  J.  L.  Hickman,  Apr 
1956  -  holotype  TM  :  K39&  ;  paratypes  I  and  4  BM(NH)  :  1973.2.53-54  ;  paratypes  2  and  8 
BJ  :  T77~78  ;  paratypes  3,  5-7  TM  :  1^397-400.  Hobart,  and  'Straham'  [possibly  Strahan, 
west  coast  of  Tasmania?]  i47°2o'E.  42°5o'S.  Map,  14/5  collector  ?  AM:  62628  -  paratypes 
9-13,  AM  :  W5339-5343. 

REMARKS.  R.  liitoralis  differs  from  all  other  species  of  the  genus  in  the  con- 
figuration of  the  male  genital  field.  The  other  Australian  species,  R.  queenslandica, 
differs  among  other  respects  in  having  only  one  pair  of  spermathecae. 


Subfamily  MEGASGOLEGINAE 
Tribe  PERIONYCHINI  Jamieson, 
Genus  PERIONYCHELLA  Michaelsen,  i907a  emend. 

DIAGNOSIS.  Setae  8  or  more  per  segment.  Combined  pores  of  vasa  deferentia 
and  tubular  or  tubuloracemose  prostates  on  XVIII.  Gizzard  in  V,  rarely  VI  or  VII 
(well  developed  to  vestigial)  ;  typhlosole  absent.  Extramural  calciferous  glands 
usually  absent,  and  never  paired,  but  the  oesophagus  often  vascularized  and  intern- 
ally rugose,  especially  in  XIV  and  XV.  Nephridia  stomate,  exonephric  holo- 
nephridia  with  or  without  bladders  ;  the  anterior  nephridia  sometimes  tufted. 
Spermathecae  I  to  5  pairs. 

DESCRIPTION.  Terrestrial  or  rarely  aquatic  worms.  Form  circular  in  cross 
section  or  depressed  ;  anus  terminal ;  with  or  without  pigmentation.  Dorsal  pores 
present  or  absent,  usually  commencing  in  3/4  or  4/5.  Setae  8  per  segment  through- 
out (lumbricine  condition)  or  becoming  more  numerous  (perichaetine  condition) 
after  a  varying  number  of  anterior  segments,  or  perichaetine  throughout.  Nephro- 
pores  in  a  single  series  throughout  in  the  vicinity  of  c  lines  (where  setae  are  lumbricine) 
or  their  equivalent  in  perichaetine  species.  Posterior  limit  of  clitellum  shortly 
anterior  to  the  male  pores  or  (P.  (P.)  canaliculatus  and  phalacrus)  including  these. 
A  pair  of  pores  of  the  combined  vasa  deferentia  and  a  pair  of  tubular,  tubuloracemose 
or  racemose  prostates  on  XVIII.  Female  pores  paired  or  exceptionally  single  mid- 
ventral,  in  XIV.  Spermathecal  pores  I  to  (usually)  5  pairs  in  front  of  or  usually 
including  intersegment  8/9.  Accessory  genital  markings  present  in  the  vicinity  of 
the  male  pores  and  sometimes  of  the  spermathecal  pores. 

Last  hearts  in  XII  or  (P.  (V.)  grandis)  XIII.  Supra-oesophageal  vessel  (always  ?) 
restricted  to  the  post-gizzard  oesophagus,  moderately  to  well  developed,  sometimes 
not  differentiated  from  the  oesophageal  plexus.  Subneural  vessel  absent.  Gizzard 
in  V  or  exceptionally  in  VI,  frequently  only  moderately  developed  or  vestigial  (in 


TASMANIAN  EARTHWORMS  217 

some  northern  Queensland  species  in  VII).  Oesophagus  commonly  with  internal 
rugae,  these  often  best  developed  in  XIV  and  XV  ;  sometimes  with  internal  laminae 
which  exceptionally  line  one  or  more  annular,  unpaired,  axial  calciferous  glands. 
Intestine  commencing  in  XVI-XX,  usually  XVII ;  typhlosole,  muscular  thicken- 
ing and  caeca  absent  or  exceptionally,  a  rudimentary  typhlosole-like  dorsal  ridge 
present.  Nephridia  simple,  exonephric  holonephridia  throughout,  or  rarely  preceded 
in  anterior  segments  by  tufted  nephridia  with  simple  or  (P.  (P.)  davallid)  composite 
ducts.  Ectal  nephridial  bladders  present  (V  esiculodrilus]  or  absent  (Perionychella)  ; 
tufted  nephridia  retaining  bladders  in  vesiculate  species.  Holandric  (testes  in  X 
and  XI),  with  seminal  vesicles  in  IX  and  XII  or  rarely  also  in  X,  or  X  and  XI,  or  in 
XI  and  XII ;  or  rarely  metandric,  with  anterior  testes  and  seminal  vesicles  reduced 
or  absent.  Metagynous  (ovaries  in  XIII)  ;  ovisacs  present  or  absent.  Prostates 
tubular,  with  central  lumen,  though  often  flattened,  or  tubuloracemose  (i.e.  lobulated 
to  externally  racemose  but  with  axial  lumen  with  (or  without  ?)  side  branches). 
Vasa  deferentia  usually  joining  the  ectal  end  of  the  prostate  gland,  occasionally  the 
junction  of  gland  and  duct.  Penial  setae  usually  present.  Spermathecae  with 
single,  clavate  or  digitiform,  uniloculate  diverticulum. 

DISTRIBUTION.  Eastern  Subregion  of  Australia  :  mostly  in  Victoria  but  extending 
northwards  to  the  Atherton  Tableland  in  North  Queensland  and  with  a  morphologic- 
ally isolated  species  near  Cooktown.  The  dominant  genus  in  Tasmania. 

REMARKS.  The  genus  Perionychella  was  established  by  Michaelsen  (i907a)  for 
four  eastern  Australian  species  :  Perichaeta  dendyi  Spencer,  i8g2b,  and  Megascolex 
pritchardi  Spencer,  1900,  both  from  Victoria ;  Perichaeta  canaliculata  Fletcher, 
i888a,  from  north  Queensland  ;  and  P.  scolecoidea  Spencer,  1895,  from  Tasmania. 
Michaelsen's  diagnosis  of  Perionychella  was  :  setae  at  least  in  the  mid-  and  hind-body 
many  (more  than  8)  per  segment.  Spermathecal  pores  2-4  (-5  ?)  pairs  between 
segments  IV  and  IX.  i  gizzard,  in  segment  5  or  6.  Completely  meganephric 
(holonephric).  Prostates  with  branched  canal  system. 

The  only  expressed  difference  from  the  typically  southwestern  Australian  Wood' 
wardia  (now  Woodwardietta  Stephenson,  1925)  was  the  increase  in  setal  numbers 
beyond  8  per  segment.  The  status  of  Woodwardiella  is  discussed  on  p.  219. 

In  1916  Michaelsen  emended  the  definition  of  the  genus  Perionyx  Perrier,  1872,  to 
agree  in  all  essentials  with  that  of  Perionychella,  which  was  therefore  suppressed,  and 
included  Diporochaeta  Beddard  in  Perionyx  as  a  subgenus.  The  subgenus  Perionyx 
was  distinguished  by  branching  of  the  prostate  duct  on  entering  the  gland  (Pheretima 
prostate)  while  in  Diporochaeta  the  gland  possessed  a  central  axial  lumen  with  or 
without  branched  or  unbranched  lateral  canals.  Perionychella  dendyi  (like  P. 
canaliculata)  was  shown  to  have  the  Diporochaeta  condition,  with  lateral  canals,  and 
was  placed  in  that  subgenus.  Stephenson  (1923)  reinstated  Diporochaeta  as  a 
separate  genus  while  suggesting  a  close  relationship  with  Perionyx.  Perionychella 
remained  suppressed  in  Diporochaeta.  Gates  (1959)  placed  Diporochaeta  and  Perionyx 
in  different  families,  the  redefined  Acanthodrilidae  and  Megascolecidae  respectively. 
Jamieson  (i97ia)  demonstrated  that  the  grounds  for  distinguishing  the  two  families 
were  invalid  and,  having  redefined  and  reconstituted  the  two  entities,  reduced  them 

13 


218  B.   G.   M.   JAMIESON 

to  subfamilial  rank  in  the  Megascolecidae.  Restriction  of  Perionyx  to  oriental 
species  (Gates,  1960)  was  accepted  and  40  Australian  species  were  provisionally 
referred  to  the  genus  Diporochaeta.  It  was  suggested  that  revision  of  Diporochaeta 
required  consideration  of  certain  species  placed  in  Plutellus  which  were  distinguished 
solely  by  the  numbers  of  setae  and  that  it  would  necessitate  establishment  of  several 
new  genera.  It  was  also  proposed  that  Perionychella  should  be  resurrected  for 
Perichaeta  dendyi  as  it  was  clearly  not  congeneric  with  the  New  Zealand  type-species 
of  Diporochaeta  (see  p.  219).  Relationship  of  Perionyx  and  Diporochaeta  s.  lat.  was 
acknowledged  by  placing  the  two  genera,  with  other  holonephric  megascolecine 
genera,  in  the  new  tribe  Perionychini. 

Revision  of  Plutellus  is  now  far  advanced  and  has  involved  resurrection  of  Argilo- 
philus  Eisen  for  American  species,  and  division  of  the  Australian  species,  after  re- 
instatement of  Fletcher odrilus  Michaelsen,  into  the  genera  Plutellus  s.  strict.,  Simsia 
Jamieson,  ig72a  ;  Graliophilus  Jamieson,  19716,  and  V esiculodrilus  Jamieson, 
I973a.  In  the  latter  paper  attention  was  drawn  to  the  close  morphological  affinity 
of  V esiculodrilus  with  similarly  vesiculate  Australian  species  of  Diporochaeta  s.  lat. 

The  Tasmanian  fauna  provides  the  key  to  revision  of  Diporochaeta  in  Australia, 
and  to  the  relationships  of  V esiculodrilus ,  as  vesiculate  perichaetine  species  which 
would  formerly  have  been  assignable  to  Diporochaeta  s.  Stephenson  are  seen  to  have 
morphological  (phenetic)  affinities,  which  are  unequivocally  intrageneric,  with  species 
attributable  to  V esiculodrilus.  Transference  of  these  perichaetine  species  to 
V esiculodrilus  greatly  reduces  the  numbers  of  species  in  Diporochaeta  but  leaves  a 
large  residue  of  species  either  lacking  nephridial  bladders  (avesiculate  species)  or 
with  the  condition  of  the  nephridial  duct  undescribed.  The  Tasmanian  fauna, 
again,  allows  determination  of  the  affinities  of  the  avesiculate  species  as  most  of 
these  on  the  island  show  high  morphological  similarities  with  V esiculodrilus.  They 
are  also  indistinguishable  from  Perionychella,  of  which  the  Victorian  type-species  is 
avesiculate.  (Variation  in  form  of  the  prostates  from  tubular  to  racemose  is  con- 
sidered unimportant  as  all  intermediates  are  seen,  author's  examination.)  It  thus 
appears  that  V esiculodrilus  must  be  included  in  Perionychella,  a  prior  genus  which 
has  been  suppressed  for  nearly  sixty  years. 

It  is  here  proposed  that  V esiculodrilus  be  retained  as  a  subgenus  of  Perionychella 
distinguished  from  the  nominate  subgenus  (P.  (Perionychella}}  in  possessing  nephridial 
vesicles.  It  is  possible  that  some  species  of  V esiculodrilus  may  be  morphologically 
closer  to  avesiculate  Perionychella  species  than  to  species  in  their  own  subgenus  (and 
vice  versa},  but  recognition  of  the  two  subgenera  achieves  one  of  the  aims  of  practical 
taxonomy,  convenience.  Separation  of  V esiculodrilus  from  Perionychella  also  finds 
some  support  in  setal  ratios  (Table  i)  though  whether  the  distinction  by  ratios  will 
be  upheld  should  additional  lumbricine  species  of  the  latter  subgenus  be  found 
remains  to  be  seen.  Thus,  for  Tasmanian  species  at  least,  dd  =  2-o-^iab>O'iS-o-22u 
in  V esiculodrilus  (with  the  exception  of  the  morphologically  very  isolated  P.  (V.) 
glandifera  in  which  dd  =  $-6ab,  o-2ju)  while  in  the  two  lumbricine  species  of  Periony- 
chella dd  =  6-5-7-8^6,  O'33-o-39w.  Furthermore,  while  recognition  of  V esiculodrilus 
species  presents  little  difficulty,  there  may  be  some  difficulty  in  ascertaining  whether 
an  avesiculate  specimen  should  be  referred  to  the  subgenus  Perionychella  or  to  other 


TASMANIAN   EARTHWORMS  219 

similarly  holonephric  genera  (particularly  the  Western  Australia  avesiculate  Gralio- 
philus)  or  merits  recognition  of  a  new  genus.  (It  should  be  noted  that  the  difficulty 
in  distinguishing  between  avesiculate  Perionychella  species  and  Graliophilus  stems 
largely  from  paucity  of  knowledge  of  some  species-groups  of  Graliophilus  and  that  it 
is  not  proposed  that  Graliophilus  should  be  united  with  Perionychella.  Rather,  as 
suggested  by  Jamieson  (19716),  Graliophilus  requires  splitting  into  further  genera.) 
Reinstatement  of  Perionychella  is  necessitated  by  the  rules  of  priority  but  Michael- 
sen's  definition  of  Perionychella  does  not  diagnose  the  majority  of  species  of  the  genus 
as  emended  and  enlarged  on  the  basis  of  consideration  of  large  numbers  of  characters, 
i.e.  overall  phenetic  resemblance  of  constituent  species,  and  an  emended  definition 
is  therefore  required.  It  will  be  shown  that  the  only  characters  which  are  constant 
throughout  the  emended  genus  are  general  characters  of  suprageneric,  often  ordinal 
or  subordinal  significance.  The  genus  is  thus  polythetic,  that  is  to  say  characters  of 
generic  significance  are  not  present  in  all  constituent  species  but  jointly  associate 
these  species  at  a  high  level  of  similarity.  Furthermore,  several  generic  characters, 
such  as  nephridial  bladders  and  five  pairs  of  spermathecae,  are  constant  for  the 
majority  of  species  and  give  striking  morphological  homogeneity  to  these  species 
which  can  often  only  be  distinguished  with  certainty  by  the  configuration  of  genital 
markings. 

Something  more  must  be  said  of  the  status  of  Woodwardiella  and  of  Diporochaeta. 
The  fact  that  in  the  present  work  Perionychella  has  been  extended  to  include  forms 
with  eight  setae  per  segment  removes  all  justification  for  separating  Woodwardiella 
as  defined  by  Michaelsen  from  this  genus.  However,  the  Western  Australian  species, 
including  the  type-species,  of  Woodwardiella  invariably  have  two  pairs  of  spermathe- 
cae and  are  characterized  by  anterior  tufted  nephridia.  These  conditions  do  occur 
exceptionally  in  Perionychella  but  it  seems  inadvisable  at  present  to  reduce  Wood- 
wardiella to  the  status  of  a  junior  synonym  in  the  former.  The  refractory  state  of 
material  of  Woodwardiella  in  the  Western  Australian  Museum  (Jamieson  1970) 
precludes  thorough  revision  of  the  genus.  The  status  of  these  Eastern  Australian 
species  tentatively  assigned,  from  poor  descriptions,  to  Woodwardiella  requires 
mention.  One  of  these,  Cryptodrilus  mortoni  Spencer,  1895,  is  shown  to  be  a 
Perionychella  (V esiculodrilus]  in  the  present  work.  A  second  C.  tesselatus  Spencer, 
1895,  is  almost  certainly  also  a  Perionychella.  It  has  not  been  rediscovered  in  this 
survey.  The  third,  Woodwardia  healesi  Michaelsen,  1923,  cannot  be  placed  with 
certainty  in  any  known  genus. 

The  type-species  of  Diporochaeta,  D.  intermedia  Beddard,  1888,  is  exceptional  in 
the  Megascolecinae  in  having  its  last  hearts  in  segment  XI  rather  than  in  XII  or 
XIII.  This  seems  a  normal  condition  in  the  species  as  it  has  been  observed  in 
material  from  several  localities  (Lee  1959).  The  fact  that  the  ovaries  are  in  the  nor- 
mal megadrile  location  in  XIII  and  that  spermathecae  may  persist  in  IX  indicates 
that  the  unusual  distribution  of  hearts  and  spermathecae  is  not  simply  due  to  deletion 
of  a  prespermathecal  metamere.  Loss  of  hearts  from  XII  would  be  contrary  to 
evolutionary  trends  in  the  Megascolecinae.  If  last  hearts  were  in  XII,  D.  intermedia 
might  be  considered  congeneric  with  the  extended  Perionychella  of  the  present  work 
and,  as  Diporochaeta  is  the  prior  genus,  all  Perionychella  species  would  have  to  be 


220  B.   G.   M.    JAMIESON 

placed  in  it.  Other  Diporochaeta  species  in  New  Zealand  appear  from  the  literature 
to  qualify  for  inclusion  in  Perionychella  s.  lat.  but  must  be  re-examined  before  this 
can  be  confirmed.  It  is  here  proposed  to  retain  Perionychella  for  Australian  forms 
conforming  with  the  Australian  type-species  Perionychella  dendyi,  and  to  allow  the 
heart-location  in  Diporochaeta  intermedia  to  merit  regarding  this  species  as  con- 
stituting a  monotypic  genus  Diporochaeta.  This  restriction  of  Diporochaeta  will  not, 
however,  be  formally  made  until  the  status  of  the  other  New  Zealand  species 
currently  in  Diporochaeta  is  decided,  when  they  may  be  referred  to  Perionychella  or 
considered  to  warrant  one  or  more  genera  distinct  from  both  Perionychella  and  a 
monotypic  Diporochaeta. 

Perionychella  accounts  for  almost  half  of  the  native  Tasmanian  earthworm 
species.  The  very  profound  emendation  of  the  genus  after  a  long  period  of  suppres- 
sion makes  it  desirable  to  list  those  non-Tasmanian  species  assignable  to  the  genus 
and  it  will  be  seen  that  these  comprise  substantial  components  of  the  fauna  of  neigh- 
bouring Victoria. 

Subgenus  PERIONYCHELLA  Michaelsen,  I9o;a  emend. 

DEFINITION.    As  for  the  genus  with  nephridia  lacking  bladders. 

DISTRIBUTION.  As  for  the  genus  but  the  generic  identity  of  the  Queensland  species 
is  uncertain. 

TYPE-SPECIES.  Perionychella  dendyi  (Spencer,  i8g2b)  Victoria.  Syntype  (?) 
examined,  AM  :  W.I2Q4. 

CHECK  LIST 
,  Tasmanian  species 

P.  (P.)  capensis  sp.  n. 

P.  (P.)  hickmani  sp.  n. 

P.  (P.)  kershawi  sp.  n. 

*Perichaeta  irregularis  Spencer,  1895 

P.  (P.)  weldboroughi  sp.  n. 

Other  species 

Unless  otherwise  stated,  material  has  been  examined  by  the  author  as  indicated. 
Victoria 
*Perichaeta   copelandi  Spencer,    i8g2b   Lectotype   NMV  :  Gi86  ;    paralectotype 

Gi443 
^Diporochaeta  davallia  Spencer,  1900,  vide  Jamieson  (1970) 

*Perichaeta  dubia  Spencer,  i892b  Syntype  (?)  AM  :  Wi293 

Megascolex   pritchardi  Spencer,    1900,   vide    Michaelsen    (i907a)    and  lectotype 

NMV  :  Gio8 
*Perichaeta  tanjilensis  Spencer,  i892b  Syntype  (?)  AM  :  Wi298 

New  South  Wales 
None  known 


TASMANIAN   EARTHWORMS  221 

Queensland 

*Perionyx  (Diporochaeta)  athertonensis  Michaelsen,  1916 

Perichaeta  canaliculata  Fletcher,  i888a,  vide  Michaelsen  (igoyb) 

*Perionyx  (Diporochaeta)  erici  Michaelsen,  1916 

*Perionyx  (Diporochaeta)  phalacrus  Michaelsen,  1916 

*Perionyx  (?)  terrareginae  Fletcher,  iSSgb 

*  Comb.  nov. 

REMARKS.  The  condition  of  the  nephridial  ducts  is  not  described  for  the  Queens- 
land species  but  they  presumably  lacked  bladders  as  all  were  described  by  Michaelsen 
(1916)  who  in  the  same  account  described  nephridial  bladders  for  a  further  Perionyx 
species  (P.  (D.)  sigillatus).  The  types  are  not  available  for  examination.  That 
these  species,  from  the  Cairns-Atherton  Tableland  area,  should  be  referred  to  the 
genus  Perionychella  is  nevertheless  questionable.  The  gizzard  is  located  more 
posteriorly  than  in  Victorian  and  Tasmanian  species,  the  number  of  setae  per  segment 
is  more  numerous  and,  although  it  is  deduced  that  nephridial  bladders  are  absent, 
they  show  affinities  with  Perionyx  (Diporochaeta)  sigillatus  Michaelsen,  1916,  from 
the  same  area,  which  has  bladders  and  appears  to  be  related  to  the  similarly  vesiculate 
genus  Fletcherodrilus,  rather  than  with  Perionychella.  Exclusion  of  the  five  north 
Queensland  species  from  Perionyx  as  emended  by  Gates  (1960)  is  necessitated  by  a 
set  of  characters  of  the  latter  genus,  viz.  female  pore  unpaired,  midventral  ;  sub- 
neural  vessel  present  ;  spermathecal  diverticula  absent. 


Subgenus  VESICULODRILUS  Jamieson, 

DEFINITION.  As  for  the  genus  with  nephridia  possessing  ectal  bladders.  (One 
morph  of  P.  (V.)  mortoni  lacks  bladders  and  their  presence  is  questionable  in  the 
type-specimens  of  P.  (V.)  lacustris.) 

DISTRIBUTION.     As  for  the  genus. 

TYPE-SPECIES.     Cryptodrilus  frenchi  Spencer,  i892a.     Victoria. 

CHECK  LIST 
Tasmanian  species 

P.  (V.)  bithecata  sp.  n. 

*Perichaeta  dilwynnia  Spencer,  1895 

P.  (V.)  evansi  sp.  n. 

P.  (V.)  glandifera  sp.  n. 

P.  (V.)  hellyeri  sp.  n. 

Cryptodrilus  hobartensis  Spencer,  1895  (new  synonym  C.  insularis  Spencer,  1895) 

*Perionyx  lacustris  Stephenson,  1924 

P.  (V.)  montisarthuri  sp.  n. 

*Cryptodrilus  mortoni  Spencer,  1895 

P.  (V.)  obliquae  sp.  n. 

P.  (V.)  pedderensis  sp.  n. 

P.  (V.)  tunnackensis  sp.  n. 

*  Comb.  nov. 


222  B.    G.   M.   JAMIESON 

Other  species 

Unless  otherwise  stated,  material  has  been  examined  by  the  author  as  indicated. 
Victoria 

*Perichaeta  alsophila  Spencer,  i892b  Syntype  (?)  AM  :  Wi297 

*Diporochaeta  arnoldi  Spencer,  1900  Lectotype  NMV  :  G203 

*Diporochaeta  euzona  Spencer,  1900  Lectotype  NMV  :  Gi2O 

Cryptodrilus  frenchi  Spencer,  18920,  vide  Jamieson  (ig73a) 

*Diporochaeta  frosti  Spencer,  1900  Lectotype  NMV  :  0117 

Cryptodrilus  gippslandicus  Spencer,  i892a,  vide  Jamieson  (1970) 

*Perichaeta  lochensis  Spencer,  i892a,  q.v. 

*Diporochaeta  manni  Spencer,  1900  Lectotype  NMV  :  Gn8 

*Diporochaeta  mediocincta  Spencer,  1900  Lectotype  NMV  :  Gi24 

*Diporochaeta  richardi  Spencer,  1900  Lectotype  NMV  :  G22O 

Cryptodrilus  tanjilensis  Spencer,  i892a,  vide  Jamieson  (i973a)  and  paralectotype 

NMV :  055 

Megascolides  tisdalli  Spencer,  1900,  vide  Jamieson  (1972)  and  lectotype  NMV  :  0178 
Plutellus  uncinatus  Stephenson,  1933,  q.v. 
Cryptodrilus  victoriae  Spencer,  18920,  vide  Jamieson  (i973a)  and  paralectotype 

NMV  :  01410 
Megascolides   volvens   Spencer,    1900,    vide   Jamieson    (ig73a)    and   syntype  (?) 

AM  :  Wi292 
*Cryptodrilus  willsiensis  Spencer,  i892a  Syntype  (?)  AM  :  Wi279 

New  South  Wales 

V esiculodrilus  frenchi  (Spencer),  vide  Jamieson  (ig73a) 
*Diporochaeta  pheretima  Jamieson,  ig73a 
V esiculodrilus  purpureus  Jamieson,  i973a 

Queensland 

*Perichaeta  barronensis  Fletcher,  i887b  Syntypes  (?)  AM  :  W2387 
*Diporochaeta  grandis  Spencer,  1900  Lectotype  NMV  :  0184 

Perionychella  species  of  uncertain  subgeneric  status 

The  following  species  which  are  not  available  for  re-examination  accord  with  the 
above-emended  definition  of  the  genus  Perionychella  but  are  of  uncertain  subgeneric 
status  as  their  authors  did  not  record  presence  or  absence  of  nephridial  bladders. 
They  are  known  customarily  to  have  omitted  reference  to  these  in  accounts  of  species 
since  shown  to  possess  bladders.  All  except  Perionychella  scolecoides  (Spencer, 
1895)  are  new  combinations  in  Perionychella. 

Tasmania 

Megascolides  bassanus  Spencer,  1895  (King  Island) 
Plutellus  decatheca  Michaelsen,  1910 
Cryptodrilus  ellisi  Spencer,  1895 


TASMANIAN  EARTHWORMS  223 

Tasmania  (cont.) 

Perichaeta  moroea  Spencer,  1895 

Perichaeta  richae  Spencer,  1895  (this  supposedly  has  gizzards  in  III  and  IV,  a 

most  doubtful  observation) 
Perichaeta  scolecoidea  Spencer,  1895 

Victoria 

Diporochaeta  apiocystis  Stephenson,  1933 
Perichaeta  bakeri  Fletcher,  i888b 
Perichaeta  dicksonia  Spencer,  i892b 
Diporochaeta  faucium  Michaelsen, 
Diporochaeta  lindti  Spencer,  1900 
Diporochaeta  nemoralis  Spencer,  1900 
Perichaeta  obscura  Spencer,  i8g2b 
Diporochaeta  sedecimalis  Michaelsen, 
Diporochaeta  spenceri  Michaelsen,  i 
Diporochaeta  telopea  Spencer,  1900 
Perichaeta  walhallae  Spencer,  i8g2b 

New  South  Wales  and  Queensland 
No  additional  species 


KEY  TO  TASMANIAN  SUBGENERA  AND  SPECIES  OF  PERIONYCHELLA 

For  species  of  uncertain  subgeneric  status  see  specific  descriptions  (p.  253  et  seq.). 

1  Nephridial  bladders  absent subgenus  PERIONYCHELLA,  2 

-  Nephridial  bladders  present       ....  subgenus  VESICULODRILUS,  6 

2  Setae  8  per  segment  throughout.*     Accessory  genital  markings  unpaired  mid  ventral 

or  paired  medianly  contiguous  respectively  in  17/18  and  18/19.     Fig.  yA 

P.  (P.)  weldboroughi  sp.  n. 
Setae  more  than  8  per  segment  throughout  or  behind  the  clitellum.     Postclitellar 

genital  markings  not  median  ..........  3 

3  Setae  8  per  segment  anteriorly  ;   numerous  posteriorly.     Fig.  56  P.    (P.)  kershawi  sp.  n. 

-  Setae  more  than  8  per  segment  throughout  .......  4 

4  Spermathecae  2  pairs.     Fig.  5A          .....  P.  (P.)  capensis  sp.  n. 

-  Spermathecae  3-5  pairs    ...........  5 

5  Spermathecae  3  pairs,  f     Fig.  6C                                          P.  (P.)  irregularis  (Spencer,  1895) 
Spermathecae  5  pairs.     Fig.  6A,  B P.  (P.)  hickmanni  sp.  n. 

6  Setae  8  per  segment  throughout          .........  7 

-  Setae  more  than  8  per  segment  throughout  or  at  least  caudally     .          .          .          .          13 

7  Spermathecal  pores  5  pairs         ..........  8 

-  Spermathecal  pores  less  than  5  pairs  .          .          .          .          .          .          .          .          10 

8  Dorsal  blood  vessel  bifid  in  the  oesophageal  region.  A  median  oesophageal  gland  in 

XV.     Seminal  vesicles  in  XI  and  XII.     Pericardiac  testis-sacs  present.     Fig. 

loA,  B  P.  (V.)  glandifera  sp.  n. 

*  An  avesiculate  morph  of  P.  (V.)  mortoni  keys  out  here  but  differs  from  weldboroughi  in  having  a 
genital  field  much  as  Fig.  I2B. 

f  Typical  specimens  of  P.  (V.)  lacustris  with  rudimentary  or  no  bladders  key  out  here  but  differ  from 
irregularis  in  having  a  genital  field  as  in  Fig.  yB. 


224 


B.    G.   M.    JAMIESON 


VI 


sp.p 


sp.p5 


Imin 


B 


FIG.  5.     Genital  fields.     A,  Perionychella  (P.)  capensis,  holotype.     B,  P.  (P.)  kershawi, 

holotype. 

Abbreviations  used  in  figures  5-32.  $,  female  pore;  g.m,  accessory  genital  marking;  £,  male  pore; 
p.s,  penial  seta;  sp.p.,  spermathecal  pore.  Roman  numerals  indicate  respective  segments.  Clitellum 
shaded.  All  figures  by  camera  lucida. 


TASMANIAN  EARTHWORMS  225 

-  Dorsal  blood  vessel  single.     No  extramural  oesophageal  glands  present  in  XV 

though  internal  rugae  may  be  present  here  and  elsehwere.     Seminal  vesicles  in 

IX  and  XII.     Testis-sacs  absent    .........  9 

9     Paired  accessory  genital  markings  of  the  male  field  at  the  posterior  borders  of  their 

segments  but  not  intersegmental.     Fig.  126    .          .       P.  (V.)  mortoni  Spencer,  1895) 

-  Paired  accessory  genital  markings  of  the  male  field  intersegmental.     Fig.  1 1 

P.  (V.)  hobartensis  (Spencer,  1895),  part 

10  Spermathecal  pores  i  pair.     Fig.  8A  .....  P.  (V.)  bithecata  sp.  n. 

-  Spermathecal  pores  2  to  4  pairs          .........         1 1 

11  Spermathecal  pores  2  pairs.     Fig.  gA.          .....       P.  (V.)  evansi  sp.  n. 

-  Spermathecal  pores  3  or  4  pairs  .........          12 

12  Spermathecal  pores  3  pairs.     Fig.  14.          .          .          .  P.  (V.)  tunnackensis  sp.  n. 

-  Spermathecal  pores  4  pairs.     Fig.  n.          .          P.  (V.)  hobartensis  (Spencer,  1895),  part 

13  Setae  8  per  segment  in  some  anterior  segments  ;  numerous  posteriorly.     Spermathe- 

cal pores  3  pairs  ............         14 

-  Setae  more  than  8  per  segment  throughout.     Spermathecal  pores  3-5  pairs    .          .          15 

14  Genital  markings  present  in  aa  in  XVII  and  XVIII.     Fig.  96.  P.  (V.)  hellyeri  sp.  n. 

-  Genital  markings  absent  in  aa.     Fig.  I2A  .          .          .  P.  (V.)  tnontisarthuri  sp.  n. 

15  Spermathecal  pores  5  pairs        ..........         16 

-  Spermathecal  pores  4  pairs  or  less       .......          .          .          17 

1 6  Accessory  genital  markings  including  a  mid  ventral  unpaired  papilla  posteriorly  in 

each  of  VI  and  VII  and  anteriorly  in  XVIII.     Fig.  8B  P.  (V.)  dilwynnia  (Spencer,  1895) 

-  Accessory  genital  markings  may  include  a  mid  ventral  papilla  in  VII  but  not  else- 

where.    Fig.  1 3 A,  B P.  (V.)  obliquae  sp.  n. 

17  Spermathecal  pores  3  pairs.     Dorsal  pores  present.     Fig.  76 

P.  (V.)  lacustris  (Stephenson,  1924) 

-  Spermathecal  pores  4  pairs.     Dorsal  pores  absent.     Fig.  126     P.  (V.)  pedderensis  sp.  n. 

Perionychella  (Perionychella)  capensis  sp.  n. 

Fig.  i,  5A,  i6A,  B  ;  Plates  91  and  92 

1  =  50  mm,  w  =  2-4,  2-5  mm,  s  =  108  (holotype  ;  paratype  i  is  a  posterior 
amputee). 

Prostomium  epilobous  f  or  proepilobous.  First  dorsal  pore  5/6.  Setae  in  XII  22, 
24 ;  caudally  31,  29  or  more  in  some  segments  ;  ventral  gap  (aa),  approximately 
constant,  =  2-^ab  throughout ;  dorsal  gap  (zz)  =  i-2yz,  narrowing  posteriorly  but 
still  distinct.  Nephropores  :  preclitellar  (XIII)  in  setal  lines  6  ;  intestinal  in  s.l.  8. 
Clitellum  annular  ^XIII-XVII.  Male  pores  on  XVIII  on  b,  each  with  a  pair  of 
protruding,  very  long  penial  setae,  on  dome-shaped  papillae  extending  from  a  to 
slightly  median  of  c.  Paired  eye-like  genital  markings  from  s.l.  i  to  s.l.  3  at  17/18 
and  from  median  to  s.l.  i  to  s.l.  2  at  19/20,  that  in  19/20  present  on  the  left  side  only 
in  paratype  i.  Female  pores  paired  anterior  and  slightly  median  of  a  on  XIV. 
Spermathecal  pores  2  pairs  on  small  papillae  in  s.l.  3  at  7/8  and  8/9. 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  in  VIII-XIII,  well 
developed.  Gizzard  vestigial,  in  V,  an  equally  large  but  thinner  walled  proventricu- 
lus  in  IV.  Extramural  calciferous  glands  absent  but  oesophagus  with  circumferential 
vascular  striae  in  VII-XV,  especially  dilated  in  XIV  and  XV  in  which,  however, 
longitudinal  rugae  remain  few  and  low.  Intestinal  origin  |XVI ;  a  very  low  dorsal 
typhlosole  present  (hardly  appreciable). 


226  B.   G.   M.    JAMIESON 

Nephridia  :  simple  stomate,  avesiculate,  exonephric  holonephridia  ;  commencing 
segment  indeterminable  ;  preseptal  funnels  demonstrated  for  those  of  XV  pos- 
teriorly. Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal 
vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  present  (paratype  i)  or 
not  developed  (holotype).  Prostates  tubuloracemose,  in  XVIII,  elongate,  tongue- 
shaped,  deeply  incised,  almost  sessile  as  the  duct  is  short ;  a  narrow  central  lumen 
present  in  the  gland.  Penial  setae  (holotype)  filiform  and  strongly  sinuous  ;  ectal 
end  in  profile  with  a  group  of  a  few  small  anteriorly  directed  spines  repeated  at 
intervals  longitudinally  ;  tip  with  long,  horn-shaped  projections  in  two  sequential 
sets  of  two,  slightly  below  the  pointed,  asymmetrical  extremity,  and  probably 
homologous  with  the  spines  ;  length  (right  setae)  1-95,  2-74  mm.  Spermathecae  2 
pairs  ;  diverticulum  (inseminated)  single,  digitiform,  uniloculate,  longer  than  the 
ampulla. 

MATERIAL  EXAMINED. 

Table  Cape,  i45°45'E.  4i°oo'S.  Map,  7/1,  Dr  J.  L.  Hickman,  24  Aug  1954  ~  holotype 
TM  :  K259  ;  paratype  i  BM(NH)  :  1972.8.3  ;  paratype  2  BJ  :  Ti. 

REMARKS.  P.  (P.)  hickmani  is  morphologically  the  closest  species  to  P.  (P.) 
capensis  in  the  subgenus.  Among  other  differences  from  hickmani  are  the  two  pairs 
of  spermathecal  pores  and  the  genital  markings  in  19/20.  It  is  also  unique  in  the 
genus,  in  Tasmania,  in  possessing  a  typhylosole-like  dorsal  thickening  of  the  intestinal 
wall  (questionably  to  be  considered  a  true  typhlosole)  and  in  location  of  the  first 
dorsal  pore  in  5/6. 


Perionychella  '(Perionychella)  hickmani  sp.  n. 

Fig.  i,  6A,  B,  i6C,  D 
1  =  94>  75  mm>  w  =  4  mm,  s  =  125,  112. 

Prostomium  epilobous  f ,  open.  First  dorsal  pore  4/5  (rudiment  in  3/4  in  holo- 
type ?).  Setae  in  XII  24,  21  ;  caudally  31,  33  ;  dorsal  and  ventral  rows  (a  and  z) 
straight  throughout ;  ventral  gap  approximately  %ab  throughout ;  dorsal  gap 
approximately  1-5  or  2yz  anterior  to  clitellum  ;  equal  toyz  caudally.  Nephropores  : 
preclitellar  pores  in  setal  lines  6-7  ;  postclitellar  in  setal  lines  7.  Clitellum  annular 
^XIII-|XVII,  XVII.  Males  pores  equatorial  in  XVIII  in  ab-b,  on  prominent 
papillae.  Paired  eye-like  genital  markings  in  16/17,  17/18  and  18/19  in  ab. 
Female  pores  paired  on  XIV,  anterior  and  slightly  median  of  a.  Spermathecal 
pores  5  pairs,  in  4/5-8/9,  on  small  papillae  in  ab. 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  in  VIII-XIII,  well 
developed.  Gizzard  large  but  compressible,  in  V.  Oesophagus  simple  ;  extramural 
calciferous  glands  absent.  Intestinal  origin  XVIII  (though  pushing  septum  17/18 
forwards)  ;  typhlosole  absent.  Nephridia  simple  stomate,  avesiculate,  exonephric 
holonephridia  commencing  in  II  (preseptal  funnel  demonstrated  in  XIV  but  pre- 
sumably present  further  anteriorly  as  nephridial  morphology  similar)  ;  tufts  absent ; 


TASMANIAN   EARTHWORMS 


227 


sp.pl 


sp.p  5 


XVI 

XVII 

XVIII 


1mm 


1mm 


FIG.  6.     Genital  fields.     A  &  B,  Perionychella  (P.)  hickmani  :   A,  holotype  ;   B,  paratype  3. 

C,  P.  (P.)  irregularis,  lectotype. 


ducts  tubular,  seminal  vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs 
large,  in  XIV.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous.  Pros- 
tates :  gland  almost  circular,  appearing  to  form  a  simple  flattened  lobe  but  resolvable 
into  a  zig-zag  with  narrow  central  lumen,  the  arms  of  which  are  expanded  and  not 
tubular  (holotype)  or  depressed  tubular,  zig-zag  in  XVII -XIX  (paratype  i)  ;  vas 
deferens  joining  ectal  end  of  gland.  Penial  setae  slender,  almost  filiform  ;  tip 
rounded,  possibly  worn,  and  with  no  notable  sculpturing  (holotype,  paratype  2)  or 
with  a  delicate  tip  hooked  through  about  sixty  degrees  (paratype  3);  length  (right 
setae),  0-96,  0-85  and  1-22  mm,  respectively.  Spermathecae  5  pairs  ;  diverticulum 
single,  clavate,  uniloculate. 

Genital  markings.  7/3  -  paratype  2  agreeing  exactly  with  holotype  and  paratype 
1  '>  I3/5  ~  paratypes  3  and  4,  mature  specimens  as  holotype  but  lacking  accessory 
genital  markings  in  18/19. 


228  B.   G.   M.    JAMIESON 

Other  variation.  The  two  mature  paratypes  from  Lake  St  Clair  agree  closely  in 
morphology  with  the  holotype,  differing  in  the  following  unimportant  respects  : 
1  =  47-49  mm  ;  s  =  105-108  ;  the  prostomium  is  closed  ;  the  setal  count  rises  to 
46  caudally  ;  the  nephridia  of  II  are  astomate  rudiments,  those  of  III  having  demon- 
strable funnels  (see  also  Genital  markings}. 

MATERIAL  EXAMINED. 

Fern  Glade,  Emu  River,  Burnie,  I45°55'E.  4i°O5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  24  Aug 
1954  -  holotype  TM  :  K26o.  Hellyer  Gorge,  i45°35'E.  4i°2o'S.  Map,  7/3,  Dr  J.  L.  Hickman, 
28  May  1954  -  paratype  J  BM(NH)  :  1972.8.12.  Parrawe,  I45°35'E.  4i°2o'S.  Map,  7/3,  Dr 
J.  L.  Hickman,  25  Aug  1954  -  paratype  2  TM  :  K26i.  Lake  St  Clair,  i46°io'E.  42°O5'S. 
Map,  13/5,  Dr  J.  W.  Evans,  Feb  1941  -paratypes  3  and  4  BM(NH)  :  1972.8.13-14;  paratype  5 
AM:  W5i85. 

REMARKS.  Differences  between  this  species  and  P.  (P.)  capensis,  which  is  closest 
to  it  morphologically,  are  given  under  the  latter  species. 

Perionychella  (Perionychella)  irregularis  (Spencer,  1895) 

Fig.  i,  6C 

Perichaeta  irregularis  Spencer,  1895  '•  53~54»  PI-  V.  fig.  52-54. 
Diporochaeta  irregularis  ;   Michaelsen,  1900  :  206  ;   Jamieson,  19715  :  83. 

1  =  87  mm,  w  =  4-7  mm,  s  =  104. 

Prostomium  canaliculate,  epilobous  |.  First  dorsal  pore  in  4/5.  Setae  n  on  each 
side  in  V,  15-16  on  each  side  in  XII  and  onto  the  midbody  ;  20  on  each  side  caudally  ; 
ventral  and  dorsal  setal  gaps  conspicuous  throughout ;  in  XII  aa  =  2ab,  zz  =  ^zy  ; 
in  the  midbody  and  caudally  aar  =  $ab,  zz  =  ^zy.  Nephropores  between  setal  lines 
8  and  9.  Clitellum  annular,  |XIII-XVII  and,  dorsally,  anterior  XVIII ;  setae, 
intersegments  (weaker)  and  dorsal  pores  retained.  Male  pores  on  prominent  elliptical 
papillae  between  setal  lines  2  and  3,  nearer  2,  on  XVIII,  the  intervening  setal  rows 
not  demonstrably  bearing  setae  in  this  segment.  Two  pairs  of  eye-like  accessory 
genital  markings  in  (or,  lectotype,  immediately  median  to)  setal  lines  3,  in  19/20 
and  20/21.  Female  pores  in  XIV,  anteromedian  of  a,  near  the  ventral  midline. 
Spermathecal  pores  3  pairs,  in  6/7,  7/8  and  8/9,  in  setal  lines  3,  4  and  5  respectively 
(Spencer)  but  only  slightly  divergent  posteriorly. 

Dorsal  blood  vessel  single,  continuous  onto  the  pharynx.  Last  hearts  in  XII 
(latero-oesophageal  but  lacking  dorsal  connectives).  Supra-oesophageal  weakly 
developed,  in  X-XII.  Gizzard  small,  with  pronounced  muscular  sheen  but  easily 
compressed,  appearing  to  lie  in  VI  but  the  very  attenuated  septum  5/6  adherent  to 
it  slightly  behind  its  anterior  limit,  suggesting  that  the  gizzard  has  moved  from  V 
into  VI  by  adhesion  of  the  attenuated  septum.  Calciferous  glands  absent  but  oeso- 
phagus vascularized  in  VIII-XVI  and  swollen  in  some  of  these.  Intestinal  origin 
XVII ;  typhlosole  absent.  Nephridia  st ornate  avesiculate  holonephridia,  commenc- 
ing in  II  (funnels  demonstrated  from  XVIII)  ;  tufted  nephridia  absent.  Holandric 
(funnels  iridescent  in  X  and  XI)  ;  seminal  vesicles  4  pairs,  small  and  simple  in  IX 
and  X,  large  and  racemose  in  XI  and  XII.  Metagynous  ;  small  ovisacs  in  XIV. 


TASMANIAN   EARTHWORMS  229 

Prostates  depressed,  linear,  laterally  deeply  incised,  in  XVII -XXII,  resolvable 
with  difficulty  into  a  tubular  form  ;  external  muscular  duct  very  long  and  tortuous. 
Penial  setae  absent.  Spermathecae  3  pairs,  not  notably  departing  from  a  line 
parallel  with  the  body  axis  ;  diverticulum  (inseminated)  single,  clavate,  uniloculate. 

TYPE-LOCALITY.   King  River  Valley,  approximately  i45°25'E.  42°io'S.  Map,  12/1. 
MATERIAL  EXAMINED. 

The  previously  dissected  lectotype,  NMV  :  Ga88. 

REMARKS.  The  above  account  considerably  augments  that  of  Spencer  and 
demonstrates  the  avesiculate  condition  of  the  nephridial  ducts  for  the  first  time. 
The  dimensions  accord  with  the  specimen  described  by  Spencer. 

Closest  to  this  species  are  the  similarly  perichaetine  species  P.  (P.)  hickmani  and 
P.  (P.)  capensis.  It  differs  from  both  in  having  three  pairs  of  spermathecal  pores 
and  in  lacking  genital  markings  anterior  to  the  male  pores. 

Perionychella  (Perionychella)  kershawi  sp.  n. 

Fig.  i,  5B,  i6E.     Table  2 
1  =  171  mm,  w  =  6-1  mm,  s  =  119. 

Prostomium  epilobous  £,  closed  with  a  grooved  peristomium  giving  a  tanylobous 
appearance.  First  dorsal  pore  4/5.  Setae  8  per  segment  anterior  to  clitellum  ; 
posterior  to  clitellum  occasionally  with  an  extra  seta  in  cd ;  in  the  posterior  half, 
setae  at  first  added  in  the  dorsal  gap,  caudally  the  rows  becoming  irregular  and 
approximately  21  seta  per  segment.  Nephropores  in  c  lines  anteriorly  ;  visible 
posteriorly  but  setal  lines  irregular.  Clitellum  annular,  XIII-XVII.  Male  pores 
on  small  dome-shaped  papillae  in  a  on  XVIII.  Small  eye-like  unpaired  midventral 
accessory  genital  markings  in  11/12  and  12/13  ;  paired  transversely  conjoined  mark- 
ings, extending  lateral  of  mid  be  at  17/18,  extending  shortly  lateral  of  b  at  18/19  and 
occupying  bb  at  19/20.  Female  pores  a  pair  in  XIV,  anteromedian  of  a.  Spermathe- 
cal pores  5  pairs  in  4/5-8/9,  on  small  papillae  in  a. 

Last  hearts  in  XII,  latero-oesophageal,  connectives  to  dorsal  vessels  very  slender, 
thread-like.  Supra-oesophageal  weak  in  VII,  well  developed  in  VIII-XII.  Gizzard 
very  large  but  easily  compressible  in  V.  Extramural  calciferous  glands  absent. 
Oesophagus  with  engorged  circumferential  vascular  striae  in  VIII -XV  ;  in  (XIII  ?), 
XIV  and  XV  strongly  dilated  and  with  longitudinal  striations  which  correspond 
with  low  internal  ridges  (severely  macerated  in  this  region)  which  suggest  some 
(calciferous  ?)  modification.  Intestinal  origin  XVII ;  typhlosole  absent.  Nephridia 
tufted  holonephridia  in  II-IX,  with  posteriorly  decreasing  numbers  of  coiled  loops, 
discharging  presetally  in  c  lines  ;  stomate  at  least  from  X  in  which  coiling  in  insuf- 
ficient to  be  termed  tufted,  but  anterior  tufted  nephridia  probably  are  also  stomate 
as  a  slender  neck  is  demonstrable  to  the  anterior  septum.  Bladders  absent  unless 
terminal  dilatations  of  the  ducts  of  the  anterior  tufts  be  considered  vestigial  bladders. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles 
racemose,  in  IX  and  XII.  Presence  of  ovaries  or  ovisacs  indeterminable  owing  to 


23° 


B.   G.   M.    JAMIESON 


maceration.  Prostates  slender  coiled  only  slightly  depressed  tubes,  winding  through 
XVII-XXIV  ;  vas  deferens  joining  the  gland  near  the  junction  of  the  sinuous 
duct.  Penial  setae  present,  form  not  elucidated  ;  length  (holotype)  2-25,  2-37, 
2-39  mm.  Spermat-hecae  5  pairs  ;  diverticulum  (inseminated)  single,  clavate, 
uniloculate. 

MATERIAL  EXAMINED. 

Under  a  rotting  log  near  the  old  Breiseis  Tin  Mine  dam,  Derby,  i47°5o'E.  4i°io'S.  Map,  9/4, 
Mr  R.  C.  Kershaw,  19  Sep  1971  -  holotype  TM  :  262. 

REMARKS.  This  species  differs  from  other  Tasmanian  species  of  the  sugbenus  in 
having  midventral  genital  markings  in  11/12  and  12/13  and  in  transition  of  the  setae 
from  8  per  segment  anteriorly  to  many  posteriorly. 


TABLE  2 

Mean  intersetal  distances  in  segment  XII  in  lumbricine  species 
of  Perionychella  (Perionychella  and  Vesiculodrilus) 


P.  (P.)  kershawi 

(i) 

12-65 

P.  (P.)  weldboroughi 

(i) 

"•57 

P.  (F.)  bithecata 

(5) 

I2-I 

P.  (F.)  evansi 

(2) 

10-35 

P.  (F.)  glandifera 

(2) 

9-89 

P.  (F.)  hellyeri 

(5) 

12-42 

P.  (F.)  hobartensis 

(12) 

n-59 

P.  (F.)  montisarthuri 

(I) 

12-5 

P.  (F.)  mortoni 

(3) 

11-83 

P.  (F.)  tunnackensis 

(!) 

11-74 

no.  of 

specimens 

aa 

P.  (P.)  kershawi 

(i) 

2-44 

P.  (P.)  weldboroughi 

(i) 

2-33 

P.  (F.)  bithecata 

(5) 

1-54 

P.  (F.)  evansi 

(2) 

1-34 

P.  (F.)  glandifera 

(2) 

2-OO 

P.  (F.)  hellyeri 

(5) 

1-52 

P.  (F.)  hobartensis 

(12) 

1-62 

P.  (F.)  montisarthuri 

(I) 

I-42 

P.  (F.)  mortoni 

(3) 

i-73 

P.  (F.)  tunnackensis 

(I) 

1-67 

no.  of 

standardized  to  a  periphery  of  100 

specimens 

aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

(i) 

12-65 

5-14 

11-46 

10-28 

33-2 

9-88 

12-25 

5-14 

(i) 

u-57 

4-96 

9-92 

9-92 

38-84 

9-92 

9-92 

4-96 

(5) 

I2-I 

7-87 

12-45 

12-76 

21-53 

13-25 

12-12 

7-87 

(2) 

10-35 

7-71 

14-28 

12-98 

20-09 

12-98 

13-9 

7-71 

(2) 

9-89 

4-94 

18-39 

8-34 

27-44 

8-18 

I7-37 

5-46 

(5) 

12-42 

8-17 

12-82 

12-51 

20-70 

12-62 

12-76 

8-00 

(12) 

n-59 

7-16 

13-22 

13-0 

21-98 

12-87 

I3-2 

6-94 

(I) 

12-5 

8-8 

ii-n 

14-81 

17-59 

15-74 

n-57 

7-87 

(3) 

11-83 

6-83 

15-89 

n-4 

19-87 

11-74 

15-79 

6-64 

(J) 

11-74 

7-04 

I3-I5 

14-08 

19-72 

14-08 

I3'I5 

7-04 

ab 

i-oo 
i-oo 

I-OO 
I-OO 
I-OO 
I-OO 
I-OO 
I-OO 
I-OO 
I-OO 


be 
2-23 

2-OO 

1-58 
1-85 


1-85 

1-26 
2-33 

1-87 


intervals/aft 
dd 

6-46 
7-83 
2-74 
2-61 

5-55 

2-53 
3-07 

2-OO 


cd 

2-OO 
2-OO 
1-62 

1-68 
1-69 

1-82 

1-68 
1-67 

2-OO 


2-9I 
2-80 


dc 
1-92 

2-OO 

1-68 
1-68 
1-66 

1-8 

1-79 

1-72 

2-OO 


cb 

2-38 

2-OO 

1-54 
i -80 

3-52 
1-56 
1-84 

2-31 

1-87 


ba 
i-oo 

I-OO 
I-OO 
I-OO 
I-IO 

0-98 

o-97 
0-89 
0-97 

I-OO 


Perionychella  (Perionychella)  weldboroughi  sp.  n. 

Fig.  i,  7A,  i6F.     Table  2 
1  =  48  mm,  w  =  1-6  mm,  s  =  71. 

Prostomium  tanylobous.     First  dorsal  pore  4/5.     Setae  8  per  segment  in  regular 
longitudinal  rows  throughout.     Nephropores  faint  in  c  lines.     Clitellum  annular 


TASMANIAN   EARTHWORMS 


231 


XIII- 


XIV- 


XVI- 


sp.p1 


XVIII 


1mm 


B 


FIG.  7.     Genital  fields.     A,  Perionychella  (P.)  weldboroughi,  holotype.     B,  P.  (V.)  lacustris, 

TM  :  K265. 

XIII-XVI.  Male  pores  equatorial  in  XVIII,  on  pronounced  papillae  in  ab  with 
centres  nearer  b  than  a  ;  preceded  by  a  pair  of  medianly  contiguous  small  papillae 
in  aa  at  17/18,  with  a  further  midventral  unpaired  papilla  at  18/19.  Faint  traces 
of  presetal  midventral  unpaired  markings  present  in  VII  ?,  VIII  ?  and  IX.  Female 
pores  a  pair  on  XIV  anterior  to  and  slightly  median  of  a.  Spermathecal  pores  5 
pairs  in  4/5-8/9  in  or  slightly  lateral  of  a. 

Last  hearts  in  XII.  Supra-oesophageal  very  weakly  developed.  Gizzard 
moderate,  but  easily  compressible,  in  V.  Calciferous  glands  absent  but  oesophagus 
greatly  dilated  in  XIII-XV  in  which  vascularization  is  increased  and  internal 
longitudinal  rugae,  though  low,  are  conspicuous.  Intestinal  origin  XVII ;  typhlosole 
absent  but  intestinal  wall  slightly  thicker  middorsally.  Nephridia  simple  stomate 
avesiculate  holonephridia  commencing  in  II  (funnels  demonstrated  in  the  intestinal 
region)  ;  ectal  ducts  lacking  bladders.  Holandric  (funnels  in  X  and  XI  iridescent)  ; 
gymnorchous  ;  seminal  vesicles  very  large,  racemose,  in  IX  and  XII.  Metagynous  ; 


232 


B.   G.   M.    JAMIESON 


1mm 


A 


sp.p1 


g.m 


B 


FIG.  8.     Genital  fields.     A,  Perionychella  (Vesiculodrilus)  bithecata,  holotype. 
B,  P.  (V.)  dilwynnia,  TM  :  Ka66. 

ovisacs  small,  in  XIV.  Prostates  flattened,  somewhat  incised,  tubular,  in  XVIII- 
XX  ;  vas  deferens  joining  gland  near  the  tortuous  duct.  Penial  setae  (holotype) 
filiform  and  sinuous,  lacking  ornamentation  but  tip  apparently  missing  ;  length 
(incomplete  ?)  =  0-42  mm.  Spermathecae  5  pairs,  diverticulum  single,  slenderly 
clavate,  uniloculate. 
MATERIAL  EXAMINED. 

1-6  miles  from  the  eastern  end  of  Weldborough  Pass,  I47°55'E.  4i°io'S.     Map,  9/3,  Dr  J.  L. 
Hickman,  26  Aug  1953  -  holotype  TM  :  K2&3- 

REMARKS.     The  midventral  genital  marking  in  18/19  diagnoses  this  species. 


TASMANIAN  EARTHWORMS  233 

Perionychella  (Vesiculodrilus)  bithecata  sp.  n. 

Fig.  i,  8A,  i6H  ;  Plate  93.     Table  2 
1  =  41,  35  mm,  w  =  1-5  mm,  s  =  93,  98. 

Prostomium  epilobous  ^  open.  First  dorsal  pore  4/5.  Setae  8  per  segment  in 
regular  longitudinal  rows  throughout.  Nephropores  in  c  lines.  Clitellum  annular 
^XIII-XVI.  Male  pores  equatorial  in  XVIII  in  ab  on  small  medianly  conjoined 
papillae  ;  2  transverse  tumid  bands  spanning  the  space  between  the  porophores,  one 
anteriorly  the  other  posteriorly  in  XVIII,  each  band  traversed  by  a  groove  which 
is  parallel  to  but  distinct  from  the  intersegmental  furrows.  Genital  markings  ;  3 
pairs  of  eye-like  markings  anterior  to  a  in  VII,  VIII  and  XVII  and  on  the  right  side 
only  in  XIX  (holotype)  ;  these  replaced  in  paratype  I  by  paired  markings  in  ab, 
posteriorly  in  VI  and  XVII.  Female  pores  a  pair  on  XIV  anteromedian  to  a  sur- 
rounded by  a  nonpigmented  oval  marking  filling  aa.  Spermathecal  pores  I  pair,  in 
6/7,  in  b. 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  moderately  de- 
veloped. Gizzard  small  to  moderate,  easily  compressible,  in  V.  Calciferous  glands 
absent ;  oesophagus  widest  in  XIV  and  XV  in  which  internally  it  has  large, 
regular  longitudinal  folds  ;  intestinal  origin  XVII  ;  typhlosole  absent.  Nephridia 
simple  stomate,  vesiculate,  exonephric  holonephridia  commencing  in  II  (funnels 
demonstrated  at  least  as  far  forward  as  the  nephridia  of  III)  ;  bladder  transversely 
ellipsoid  and  projecting  median  of  the  pore  (in  c  line)  through  not  truly  diverticulate. 
Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles  race- 
mose in  IX  and  XII.  Metagynous  ;  ovisacs  small  in  XIV.  Prostates  depressed 
tubular,  compacted  coils  in  XVI I -XXI ;  Penial  setae  (paratypes  i,  3)  filiform  and 
sinuous  ;  tip  inrolled,  hooked  and  with  a  pair  of  'dorsal',  short  horns  visible  only 
under  the  electron  microscope  ;  length  (paratype  i)  0-39  mm.  Spermathecae  i  pair, 
with  pores  in  6/7  ;  the  ampulla  extending  into  IX  or  VIII ;  diverticulum  (insemi- 
nated) single,  uniloculate,  bent  clavate. 

Genital  markings  in  the  holotype  and  3  paratypes 

Total  number 
Specimen  of  specimens 

Paired  posterior  in  VI  in  ab  -  Pi.     P3(R)  2 

Paired  presetal  in  VII  in  a  H  - 

Paired  presetal  in  VIII  in  a  H  - 

Paired  presetal  in  XVII  in  a  H  - 

Paired  posterior  in  XVII  in  ab  -  Pi,  2.     P3(R) 

Unilateral  (R)  in  18/19  in  ab  -  P2 

Unilateral  presetal  in  XIX  in  a  (R)  H  — 

or  ab  (L)  -  P3 
H  =  Holotype;  P  =  Paratypes.     R  =  right.    L  =  left. 

MATERIAL  EXAMINED. 

Hellyer  Gorge,  I45°35'E.  4i°2o'S.  Map,  7/3,  Dr  J.  L.  Hickman,  28  May  1954  -  holotype 
TM  1X264;  paratype  i  BM(NH)  :  1972.8.1  ;  paratypes  2,  3  AM  :  W$  186-7  :  paratype  4 
BJ  :  T2. 

14 


234  B-   G-  M-   JAMIESON 

REMARKS.  The  configuration  of  genital  markings  in  the  paratypes  differs 
markedly  from  that  in  the  holotype  but  similarity  of  general  morphology,  including 
the  bithecal  condition  (which  is  unique  in  the  genus),  indicates  conspecificity. 


Perionychella  (Vesiculodrilus)  dilwynnia  (Spencer,  1895) 
Fig.  i,  8B,  i6J  ;  Plates  94-96 

Perichaeta  dilwynnia  Spencer,  1895  :  50-51,  PI.  IV,  fig.  46-48. 
Diporochaeta  dilwynnia  ;   Michaelsen,  1900  :  204  ;   Jamieson  igyib  :  83. 

1  =  71,  52  mm,  w  =  3-0,  2-6  mm,  s  =  104,  118  (Specimens  i  and  2  respectively, 
throughout  account). 

Prostomium  epilobous  £,  open,  but  deep  dorsal  canalicula  extending  to  inter- 
segment 1/2.  Peristomium  similarly  grooved  midventrally.  First  dorsal  pore  4/5. 
Caudal  end  dorsoventrally  depressed.  Setae  in  XII,  16,  14  ;  caudally  24  per  seg- 
ment ;  dorsal  and  ventral  rows  (a  and  z)  straight  throughout  but  caudally  some  setae 
of  z  line  are  omitted  ;  ventral  gap  approximately  2ab  throughout ;  dorsal  gap  in- 
constant even  over  a  few  segments,  usually  a  definite  interruption  of  the  setal  circlet 
but  not  more  than  twice  the  general  zy  width.  Nephropores  :  preclitellar  in  setal 
lines  4-5  ;  postclitellar  in  4  ;  caudally  in  6-7  ;  in  a  straight  lateral  row  on  each  side. 
Clitellum  annular,  XIII,  £XIII-XVII.  Male  pores  equatorial  in  XVIII  in  mid  ab, 
on  large  prominent  papillae  ;  a  transversely  elongate  elliptical  papilla  filling  the 
space  between  them  in  the  anterior  half  of  the  segment.  Paired  eye-like  genital 
markings  in  19/20  and  20/21  in  ab  and  an  unpaired  postsetal  midventral  marking  in 
each  of  VI  and  VII.  Female  pores  shortly  anteromedian  to  setae  a  of  XIV.  Sperma- 
thecal  pores  5  pairs,  on  small  papillae  in  4/5-8/9,  in  ab. 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  well  developed  in 
IX-XII,  weak  in  VIII  and  XIII.  Gizzard  small,  almost  rudimentary,  in  V. 
Oesophagus  simple,  extramural  calciferous  glands  absent.  Intestinal  origin  XVII ; 
typhlosole  absent.  Nephridia  simple  stomate,  exonephric,  vesiculate  holonephridia 
throughout,  commencing  in  II ;  each  with  large,  single  preseptal  funnel  and  long, 
wide,  tortuous  tubular  bladder.  Holandric  (funnels  iridescent)  ;  gymnorchous  ; 
seminal  vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  in  XI V.  Prostates 
depressed,  lobulated  tubular,  with  compressed  coils,  in  XVII-XX  (i),  XVIII  only 
(2)  ;  vas  deferens  joining  junction  of  gland  and  duct.  Penial  setae  (specimens  i 
and  2)  slender,  needle-like,  not  filiform  ;  tip  pointed,  asymmetrical,  convex  on  one 
side,  straight  or  slightly  concave  on  the  other  ;  ornamentation  in  profile  consisting 
of  alternating  V-shaped  depressions  the  border  of  which,  at  the  angle,  projects 
towards  the  apex  of  the  seta  as  an  irregular  scale  ;  length  (specimen  i)  =  0-77  mm. 
Spermathecae  5  pairs  ;  diverticulum  (inseminated)  single,  clavate,  uniloculate. 

Genital  markings.  13/4  -  (specimens  1-9)  median  genital  marking  in  VI  absent  in 
specimens  3-6,  8  ;  otherwise  agreeing  in  all  respects  with  specimen  i. 

TYPE-LOCALITY.     Dee  Bridge,  i46°35'E.  42°i5'S.     Map,  13/7. 


TASMANIAN  EARTHWORMS 


235 


g.m 


VI 


XVI 


1mm 


B 


FIG.  9.     Genital  fields.     A,  Perionychella  (Vesiculodyilus)  evansi,  holotype. 
B,  F.  (V.)  hellyeri,  holotype. 


236  B.  G.  M.  JAMIESON 

MATERIAL  EXAMINED. 

Tarraleah,  i46°25'E.  42°2o'S.  Map,  13/4,  over  pipeline,  Dr  J.  L.  Hickman,  27  May  1954  ~ 
specimens  1-9  TM  :  K266-274  ;  specimens  10-13  BM(NH)  :  1972.8.4-7  ;  Butlers  Gorge  Road, 
I -4  miles  from  Tarraleah,  Dr  J.  L.  Hickman,  27  May  1954  ~  specimens  14-20  TM  :  1^275-281  ; 
specimens  21-32  BJ  :  T3- 14  ;  Tarraleah,  Lyell  Highway,  Dr  J.  L.  Hickman,  22  May  1954- 
specimens  33-36  AM  :  W5i88~5i9i. 

REMARKS.  The  new  material  was  collected  a  few  miles  from  the  Dee  Bridge 
type-locality.  The  type-specimens  are  lost  (Jensz  and  Smith  1969).  It  appears 
from  Spencer's  description  that  they  differed  from  the  Tarraleah  specimens  in  clitel- 
lar  extent  (XIV-XVII)  ;  in  location  of  the  paired  genital  markings  in  18/19  and  19/20 
and  absence  of  the  midventral  markings  in  VI  and  VII ;  the  median  marking  in 
XVIII  was  present  and  setal  counts  agreed  closely.  Nephridial  bladders  were  not 
mentioned. 

Perionychella  (Vesiculodrilus)  evansi  sp.  n. 

Fig.  i,  gA,  156,  i6K.    Table  2 
1  =  51  (?)  mm  (the  paratype  damaged  anteriorly),  w  =  2-6,  2-5  mm,  s  =  102, 117. 

Prostomium  epilobous  §,  closed  ;  first  dorsal  pore  (2/3  ?),  3/4  (holotype).  Setae  in 
8  regular  longitudinal  rows  throughout.  Nephropores  in  c  lines.  Clitellum  annular, 
XIII -XVI.  Male  pores  on  small  papillae  in  a  on  XVIII.  Five  transversely  ellip- 
tical, unpaired,  midventral  genital  markings  in  the  anterior  half  of  VII,  VIII,  IX, 
XVIII  and  XIX  (holotype),  that  in  IX  absent  in  paratype,  and  a  pair  of  eye-like 
markings  at  18/19  with  centres  in  b  (both  specimens).  Female  pores  a  pair  in  XIV 
anteromedian  of  a.  Spermathecal  pores  2  pairs  in  7/8  and  8/9,  in  a. 

Last  hearts  in  XII  (laterb-oesophageal).  Supra-oesophageal  well  developed. 
Gizzard  in  V  (damaged  and  not  recognizable  in  paratype),  moderately  large  but  fairly 
easily  compressible.  Calciferous  glands  absent  but  oesophagus  internally  rugose  in 
VI -XV,  especially  so  in  XIV  and  XV  in  which  it  is  conspicuously  dilated,  but  lacking 
specialized  internal  lamellae.  Intestinal  origin  XVI ;  typhlosole  absent.  Nephridia 
simple  stomate  vesiculate  exonephric  holonephridia  ;  first  seen  in  III  but  pores 
apparently  commencing  anteriorly  in  II ;  funnels  present  at  least  as  far  forward  as 
VII  ;  the  bladders  at  first  pyriform  but  by  IX  becoming  distinctly  bilobed,  the  ectal 
nephridial  duct  joining  the  median  lobe.  Holandric  (funnels  very  large,  only  those 
in  X  iridescent  in  the  holotype,  iridescent  in  XI  also  in  the  paratype)  ;  seminal 
vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  very  large,  multiloculate, 
in  XIV  (holotype  ;  not  demonstrable  in  paratype) .  Prostates  appearing  racemose, 
in  XVII-XX,  XXI  but  resolvable  into  a  compressed  and  minutely  lobulated  tubular 
form.  Penial  setae  (holotype)  needle  like,  not  filiform  ;  tip  smooth  and  rounded  ; 
ornamentation  absent,  length  =  0-71  mm.  Spermathecae  2  pairs  ;  diverticulum 
(inseminated)  single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

Lake  St  Clair,  i46°io'E.  42°O5'S.  Map,  13/5,  Dr  J.  W.  Evans,  Feb  1941  -  holotype  TM  : 
K282.  Same  locality,  Tasmanian  Biological  Survey  :  J 20,  Feb  1941 -paratype  BM(NH)  : 
1972.8.8. 


TASMANIAN  EARTHWORMS 


237 


sp.p  5 


1mm 


FIG.  10. 


1mm  D 

Genital  fields.    Perionychella  ( Vesiculodrilus)  glandifera  :  A,  holotype  ; 
B,  paratype. 


REMARKS.  This  species  appears  to  be  closest  morphologically  to  P.  (V.)  lacustris. 
Some  differences  from  the  latter  are  the  lumbricine  setae,  the  two  pairs  of  spermathe- 
cal  pores,  the  presetal  rather  than  postsetal  location  of  the  anterior  genital  markings 
and  the  configuration  of  those  of  the  male  genital  field. 


Perionychella  (Vesiculodrilus)  glandifera  sp.  n. 

Fig.  i,  loA,  B,  150,  i6L,  M.     Table  2 
1  =  75,  73  mm,  w  =  6-5,  7-5  mm,  s  =  98. 

Prostomium  canaliculate,  broadly  tanylobous.  Body  rectangular  in  cross  section. 
First  dorsal  pore  3/4  (imperf orate  ?  holotype),  4/5  perforate.  Setae  8  per  segment  in 
regular  longitudinal  rows  throughout.  Nephropores  in  c  lines.  Clitellum  annular, 
XIII- £X VIII  (holotype  ;  the  paratype  is  aclitellate) .  Male  pores  equatorial  in  a  lines 
of  XVIII  on  weakly  developed  papillae  on  the  setal  ridge  ;  2  pairs  of  large  sucker-like 
genital  markings  with  pore-like  centres  in  intersegments  17/18  and  18/19,  the  centres 
lateral  to  b  lines,  each  marking  spanning  the  distance  between  the  setal  arcs  of  ad- 
jacent segments  ;  a  transverse  glandular  depression  linking  each  marking  with  that 


238  B.   G.   M.   JAMIESON 

of  the  other  side  (holotype).  Two  pairs  of  eye-like  transversely  conjoined  markings 
shortly  lateral  of  b  at  17/18  and  18/19  m  paratype.  Female  pores  shortly  anterior 
to  the  setal  arc  of  XIV  at  1/3  aa.  Spermathecal  pores  minute,  5  pairs  in  4/5-8/9  in  a 
lines. 

Dorsal  vessel  segmentally  bifid  in  VIII  (paratype),  IX  (holotype) -XVIII ;  last 
hearts  in  XI I  (latero-oesophageal).  Supra-oesophageal  weakly  developed.  Gizzard 
large,  in  V  but  extending  well  posteriorly.  Oesophagus  in  XV  expanded  to  form  a 
subspherical  unpaired  calciferous  gland  with  many  closely  packed  internal  radial 
laminae  which  fill  the  oesophageal  lumen  but  do  not  unite  centrally.  Intestinal 
origin  ^XVII  (paratype)  or  posterior  XVII  (holotype)  ;  typhlosole  absent.  Nephri- 
dia  vesiculate,  exonephric  holonephridia  discharging  by  large  bladders  throughout, 
commencing  in  II  (holotype)  ;  those  in  II  forming  small  tufts  and  lacking  detectable 
funnels  (no  nephridia  detectable  in  II  in  paratype)  ;  the  remaining  nephridia  simple 
and  each  with  a  preseptal  funnel ;  bladders  in  the  anterior  segments  very  long,  wide 
tortuous  tubes  ;  in  VI  (holotype)  or  somewhat  more  posteriorly  (paratype)  the  blad- 
der is  bent  near  its  ectal  end  and  the  bend  is  drawn  out  slightly  as  a  diverticulum  ; 
succeeding  bladders  are  increasingly  diverticulate  ;  full  development  of  the  diverti- 
culum as  a  long  lateral  digitiform  process  is  attained  by  segment  XIX  and  is  main- 
tained to  the  caudal  extremity  although  caudal  nephridia  are  smaller  than  those  in 
the  anterior  region.  Holandric  (funnels  iridescent)  ;  cleistorchous,  pericardiac 
testis-sacs  formed  by  longitudinal  fusion  of  septa  9/10,  10/11  and  11/12  ;  seminal 
vesicles  racemose  in  XI  (in  the  testis-sac)  and  XII  (free).  Metagynous.  Ovisacs 
absent  (holotype)  or  present  in  XIV  (paratype).  Prostates  depressed  tubular, 
compactly  coiled,  in  XVIII-XX  (holotype)  ;  restricted  to  XVIII  and  not  fully 
developed  in  paratype  ;  vas  deferens  joining  ectal  end  of  gland  (holotype)  ;  penial 
setae  absent.  Spermathecae  5 'pairs  ;  diverticulum  (inseminated  only  in  holotype) 
single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

St  Helens,  on  road  to  Launceston  via  Scottsdale  between  94  and  96  mile  posts,  I48°I5'E. 
4i°2o'S.  Map,  10/2,  Dr  J.  L.  Hickman,  26  Aug  1953  -  holotype  TM  :  K283.  St  Columba  Falls, 
I47°55'E.  4i°2o'S.  Map,  9/2,  Dr  J.  L.  Hickman,  17  Apr  1954  -  paratype  BM(NH)  :  1972.8.9. 

REMARKS.  Perionychella  (V.)  glandifera  is  unique  among  the  known  Tasmanian 
species  of  the  genus  in  the  bifid  dorsal  vessel,  the  extramural  oesophageal  gland  in 
XV,  in  possessing  testis-sacs  and  in  location  of  the  seminal  vesicles  in  XI  and  XII. 


Perionychella  (Vesiculodrilus)  hellyeri  sp.  n. 

Fig.  i,  96,  i6N,  O.     Table  2 
1  =  54,  62  mm,  w  =  2-6,  3-0  mm,  s  =  104,  97. 

Prostomium  indistinctly  epilobous  |,  open.  First  dorsal  pore  4/5.  Setae  8  per 
segment,  the  rows  becoming  irregular  in  the  posterior  12  or  40  segments  in  which 
there  are  20-24  setae  per  segment.  Nephropores  in  c  lines.  Clitellum  annular, 
£XIII-XVI.  Male  pores  on  prominent  rounded  papillae,  equatorial  in  XVIII,  in  b  ; 


TASMANIAN  EARTHWORMS  239 

the  papillae  transversely  conjoined  by  a  midventral  boss  which  bears  presetally  two 
widely  conjoined  pore-like  markings,  a  similar  boss  present  with  conjoined  presetal 
markings  on  XVII ;  a  pair  of  eye-like  genital  markings  present  in  17/18  in  b  lines 
(holotype)  ;  presetal  markings  absent  from  XVII  in  paratype  I  and  the  median 
boss  in  XVIII  replaced  with  2  small  circular  markings  in  aa  joined  to  the  male 
papillae  by  slight  ridges  ;  eye-like  markings  present  in  17/18.  Two  pairs  of  eye-like 
markings  presetally  in  b,  on  IX  and  X  in  paratype  I  ;  absent  in  holotype.  Female 
pores  paired,  on  XIV,  anteromedian  of  a.  Spermathecal  pores  3  pairs,  in  6/7-8/9, 
in  b,  on  small  papillae  each  of  which  is  preceded  by  a  lip-like  prominence  (holotype, 
paratypes  1-5). 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  in  VIII-XIII,  well 
developed.  Gizzard  vestigial  (questionably  present),  in  V.  Extramural  calciferous 
glands  absent  but  oesophagus  enlarged  and  vascularized  with  lamellar  internal  rugae 
in  XIV  and  XV.  Intestinal  origin  XVII ;  typhlosole  absent.  Nephridia  simple, 
vesiculate  exonephric  holonephridia ;  preseptal  funnel  demonstrated  in  those  of 
III ;  possibly  rudimentary  nephridia  in  II ;  bladder  large,  subspherical  to  trans- 
versely elliptical,  the  lateral  extremity  protuberant  and  almost  forming  a  diverti- 
culum.  Holandric  (gymnorchous)  or  probably  incipiently  metandric  ;  only  posterior 
funnels  iridescent  and  seminal  vesicles  large  in  XII  but  vestigial  in  IX  (holotype)  ; 
male  organs  of  IX  and  X  completely  absent  in  paratypes  I  and  2  (metandric). 
Metagynous  ;  ovisacs  in  XIV.  Prostates  flattened  tubular,  with  lobulated,  com- 
pacted coils  in  XVIII-XX,  XXI.  Penial  setae  (paratype  i)  moderately  stout,  not 
filiform,  but  ectal  region  sigmoid ;  tip  simple  ;  ectally  weakly  ornamented  with 
anteriorly  directed  ^-shaped  scales  flush  with  the  surface  and  commonly  in  triads  ; 
the  tip  of  a  reserve  seta  (paratype  5)  has  a  delicate,  hooked  extremity  ;  length  of  a 
well-developed  seta  (paratypes  2  and  5)  =  1-18,  i-oo  mm.  Spermathecae  3  pairs  ; 
diverticulum  (inseminated)  single,  clavate,  uniloculate. 

Genital  markings  of  the  holotype  and  5  paratypes 

Total  number 
Specimen  of  specimens 

Paired  presetal  in  a  of  VII  —  Pi,  5  2 

Paired  presetal  in  a  of  VIII  -  PS  i 

Paired  presetal  in  a  of  IX  -  PS,  5  2 

Paired  presetal  in  b  of  IX  -  Pi  i 

Paired  presetal  in  i  of  X  -  Pi  I 

Paired  presetal  in  aa  of  XVII  H  Pa,  3,  5  4 

Paired  presetal  in  aa  of  XVIII  H  Pi -5  6 

Paired  in  b  in  17/18  H  Pi -5  6 
H  =  Holotype  ;   P  =  Paratypes . 

Spermathecal  pores  and  male  porophores  invariable  in  6  specimens. 
MATERIAL  EXAMINED. 

Hellyer  Gorge,  I45°35'E.  4i°2o'S.  Map,  7/3,  Dr  J.  L.  Hickman,  28  May  1954  -  holotype 
TM  :  Ka84  ;  paratypes  i  and  2  BM(NH)  :  1972.8.10-11  ;  paratypes  3  and  4  AM  :  W5I92- 
5193  ;  paratype  5  BJ  :  115.  Mt  Arthur,  in  rainforest,  i47°2o'E.  4i°i5'S.  Map,  9/1,  Mr  A.  J. 
Dartnall,  13  Mar  1971  -  2  specimens  TM  :  1^285-286. 


240 


B.   G.   M.   JAMIESON 


XIV 


XVI- 


sp.pl 


sp.p  5 


sp.p  1 


sp.p  5 


sp.p 


sp.p  5 


1mm 


1mm 


TASMANIAN  EARTHWORMS  241 

REMARKS.    This  species  is  close  morphologically  to  P.  (V.)  montisarthuri  but  is 
readily  distinguished  from  it  by  the  presence  of  postclitellar  genital  markings  in  aa. 


Perionychella  (Vesiculodrilus)  hobartensis  (Spencer,  1895) 
Fig.  i,  nA-F,  I5A,  B,  i6P-R  ;  Plates  97,  98.    Table  2 

Cryptodrilus  hobartensis  Spencer,  1895  :  37-38,  PL  I,  fig.  10,  u,  12. 
Plutellus  hobartensis  ;  Michaelsen,  1900  :  175-176  ;   Jamieson,  19710  :  87. 
Cryptodrilus  insularis  Spencer,  1895  :  41-42.     PL  2  ;  fig.  19-21  (new  synonym). 
Plutellus  insularis  ;  Michaelsen,  1900  :  176  ;   Jamieson,  19710  :  87. 
Vesiculodrilus  hobartensis  ;   Jamieson,  I973a  :  225. 
Vesiculodrilus  insularis  ;   Jamieson,  i973a  :  225. 

1  =  28  mm+  (posterior  regenerate),  w  =  2-7  mm,  s  =  91  +  . 

Form  moderately  stout ;  approximately  circular  in  cross  section  but  flattened 
between  adjacent  setae.  Prostomium  epilobous  f,  open  but  apparently  with  a 
transverse  groove  at  about  half  peristomium  ;  a  middorsal  groove  commencing  on 
the  prostomium  and  continuous  throughout  the  body.  First  dorsal  pore  4/5.  Setae 
readily  visible,  in  8  regular  longitudinal  rows  throughout,  commencing  on  II ;  ab 
absent,  cd  present,  on  XVIII. 

Nephropores  in  c  lines  at  the  anterior  borders  of  their  segments  (clitellum  and 
posteriorly).  Clitellum  annular,  strongly  protuberant  on  XIV-XVI  but  XVII  with 
clitellar  modification,  especially  dorsally  over  the  anterior  third  ;  dorsal  pores  present 
but  imperforate  ;  setae  and  intersegmental  furrows  retained  but  less  distinct ; 
nephropores  visible.  Small  indistinct  (imperforate)  male  porophores  in  ab  nearer 
b,  of  XVIII ;  paired  eye-like  accessory  genital  markings  in  a  lines  in  16/17, 17/18  and 
18/19,  the  anterior  pair  large  and  more  conspicuous  than  the  second  pair,  the  pos- 
terior pair  rudimentary.  Female  pores  minute,  shortly  anteromedian  of  a  in  XIV, 
each  with  a  distinctly  visible  white  'halo'.  Spermathecal  pores  not  visible. 

Last  hearts  in  XII  (latero-oesophageal)  ;  connections  of  the  4  pairs  of  hearts  with 
the  dorsal  vessel  presumably  present  but  not  verified  with  certainty  as  hearts  were 
brittle  and  were  separate  from  the  dorsal  vessel  on  examination.  Supra-oesophageal 
well  developed  in  VIII-XII,  and  with  a  slender  posterior  continuation  into  XIII  and 
apparently  intramurally  to  |XV. 

Gizzard  in  V,  almost  cylindrical  and  only  slightly  wider  than  the  oesophagus  but 
recognizable  by  its  muscular  sheen  ;  easily  compressible  ;  preceded  in  IV  by  a  less 
muscular  proventriculus  of  the  same  width,  from  which  it  is  not  separated  by  any 
appreciable  constriction.  Oesophagus  narrow  (though  not  much  narrower  than  the 
gizzard)  and  not  evidently  vascularized  in  VI  and  VII ;  still  narrow  but  with  intra- 
mural circumferential  vascular  striae,  which  form  the  supra-oesophageal  vessel  in 
VIII-XI ;  forming  a  segmental  unpaired  annular  dilatation  in  each  of  segments  XII, 


FIG.    u.     Genital  fields.     Perionychella   (Vesiculodrilus)   hobartensis  :    A,   lectotype ;     B, 
13/6,  TM  :  K293  ;    C,  14/2,  TM  :  K299  ;    D,  14/1,  TM  :  K294  ;    E,  19/2,  AM  : 
F,  14/6,  TM  :  K3o6. 


242  B.   G.   M.    JAMIESON 

XIII  and  XIV,  the  dilatations  increasing  in  size  posteriad ;  the  oesophagus  also 
widened,  but  less  so,  to  ^XV  but  narrow  and  chloragenous  looking  in  the  posterior 
half  of  this  segment ;  the  internal  walls  of  the  dilatations  with  rounded  longitudinal 
ridges  too  low  to  be  considered  lamellae.  Intestinal  origin  XVII ;  typhlosole 
absent. 

Genital  markings 

Specimen  Paired  genital  markings 

15/16       16/17       17/18       18/19       19/20       20/21 

Spencer,  1895  +  +  +  + 

Lectotype  +  +  +  ?  + 

7/2  TM :  K288  +  +  +  + 

8/2  TM :  K28g  +  R 

8/2  TM :  K290  +  + 

10/1  TM :  K2Qi  +  +  + 

10/1  TM :  K292  +  +  + 

13/6  TM :  K293  + 

14/1  TM :  K294  R  + 

14/1  TM :  K295  + 

14/1  TM :  K296  + 

14/1  TM :  K297  + 

14/2  TM :  K299  +  +  + 

14/3  TM :  K3oo  +  +  +  ? 

14/4  TM :  K302  +  L  +  + 

14/5  Lenah  V  TM :  K304  +  +  +  + 

14/5  Sandy  Bay  TM :  K305  +  +  +  +  R 

14/6  Risdon  TM :  K3o6  +  + 


14/6  E.  Risdon     BM(NH) 
14/6  BM(NH) 

14/6  BM(NH) 


1972.8.15  +  +  + 

1972-8.16  +  +  + 

1972.8.17  +  +  + 


19/2  AM :  W5I94  +  +  R 

19/2  AM :  W5I95  +  + 

19/3  TM :  K3o8  +  + 

L  =  left  only ;  R  =  right  only. 

Nephridia  simple  vesiculate  holonephridia  throughout ;  the  first  discharging  at 
intersegment  2/3,  each  with  a  wide  elongate  bladder  bent  into  a  U  or  J  shape,  all 
bladders  discharging  in  front  of  setae  c  ;  'necks'  to  septa  seen  but  no  funnels  demon- 
trated  with  certainty.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  seminal  vesicles 
not  recognizable.  Prostates  represented  only  by  thick  muscular  ducts  each  with  the 
form  of  a  short,  straight,  entally  narrowing  cone.  Penial  setae  not  detectable  in  the 
lectotype  and  not  reported  by  Spencer  (1895)  ;  demonstrated  in  one  or  (loc.  19/2)  two 
specimens  from  each  of  localities  7/2,  8/2,  10/1,  14/1,  14/5,  14/6  and  19/2  ;  filiform 
and  sinuous,  tip  frequently  worn  or  damaged  but  intact  in  14/5  and  14/6  in  which  it 
is  widely  bifid,  while  simple  and  hooked  in  7/2  ;  ornamentation  a  few  faint  longi- 
tudinal ridges  (14/5,  14/6)  or  groups  of  several  scarcely  protuberant,  approximately 
palmately  arranged,  apically  directed  teeth  at  intervals  near  the  tip  (7/2)  or  a  few 
very  faint  triangular  marks  and  longitudinal  ridges  (14/6)  ;  length  =  0-97,  1-56  mm 
(19/2),  1-71  mm  (14/6).  Ovaries  (laminate  with  several  strings  of  large  oocytes)  and 


TASMANIAN  EARTHWORMS  243 

stout  funnels  in  XIII  ;  ovisacs  absent.  Spermathecae  5  pairs,  the  last  in  IX  ;  with 
ovoid  ampulla  sessile  on  the  body  wall  and  a  lateral  clavate  diverticulum  as  long  as 
or  longer  than  the  ampulla.  In  one  spermatheca  the  diverticulum  is  very  short 
and  there  is  a  further  diverticulum  on  the  median  aspect  of  the  spermatheca  further 
entally.  Length  of  the  right  spermatheca  of  IX  =  0-75  mm  ;  ratio  total  length  : 
length  diverticulum  =  0-8. 

Other  variations.  Spermathecal  pores  are  4  pairs  in  the  material  listed  above 
from  localities  7/2  and  14/1,  but  5  pairs  in  all  other  specimens  examined.  The  gizzard 
is  very  large  in  specimens  from  locality  10/1,  small  to  vestigial  in  all  others.  In- 
testinal origin  is  XVII  (Spencer  1895  ;  lectotype  ;  localities  8/2,  10/1,  13/6,  14/2, 
14/3,  14/4,  14/5,  14/6)  ;  JXVII  (locality  14/1)  ;  XVI  (locality  14/6,  19/2)  or  |XVI 
(locality  7/2) .  Penial  setae  are  present  in  all  specimens  examined  with  the  exception 
of  the  lectotype.  The  latter  specimen  lacks  seminal  vesicles,  has  prostate  ducts 
but  no  glands  and  has  abnormal  Spermathecae.  These  features  suggest  that  it  is  a 
parthenogenetic,  uniparental  morph  and  that  absence  of  penial  setae,  in  a  species 
normally  possessing  these,  is  further  evidence  of  parthenogenesis. 

Ovisacs  were  demonstrated  for  specimens  from  all  localities  excepting  13/6  and 
14/5  and  the  lectotype. 

TYPE-LOCALITIES.     Parattah  and  Mt  Wellington. 
MATERIAL  EXAMINED. 

Parattah,  I47°25'E.  42°2o'S.  Map,  14/1,  Baldwin  Spencer  collection,  Jan  1893 -the  pre- 
viously undissected,  clitellate  lectotype,  in  poor  condition,  NMV  :  050.  The  two  paralecto- 
types,  051,  were  examined  but  were  not  used  for  the  above  account  as  one  is  broken  in  two  at  the 
clitellum  and  the  other  is  imperfectly  clitellate. 

The  following  new  material  has  been  identified  on  external  and  internal  anatomy  :  Fern  Glade, 
Emu  River,  I45°55'E.  4i°o5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  24  Aug  1954 -TM  :  K28y-288. 
Great  Lake,  near  stones  around  base  of  gum  tree,  slopes  of  lake,  i46°45'E.  4i°55'S.  Map,  8/2, 
Dr  J.  L.  Hickman,  26  May  1954  ~  TM  :  1^289-290.  Goulds  County,  near  Lottah,  fireplace  at 
creek,  I48°05'E.  4i°i5'S.  Map  10/1,  Dr  J.  L.  Hickman,  16  Apr  1954 -TM :  K2gi -292. 
Marlborough  Highway,  near  Bronte,  i46°3o'E.  42°io'S.  Map,  13/6,  Dr  J.  L.  Hickman,  26 
May  I954-TM  :  K293.  Parattah  (type-locality)  I47°25'E.  42°2o'S.  Map,  14/1,  under  moss, 
in  earth  at  base  of  cliff,  also  in  earth  along  sides  of  logs.  Map,  14/1,  Professor  V.  V.  and  Dr  J.  L. 
Hickman,  18  Aug  1954 -TM  :  1^294-298.  Tunnack,  under  logs  and  stones,  damp  conditions, 
i47°3o'E.  42°25'S.  Map,  14/2,  Dr  J.  L.  Hickman,  18  Aug  1954  -  ™  :  K299.  Collinsvale  near 
Hobart,  Myrtle  Forest,  I47°O5'E.  42°5o'S.  Map,  14/3,  Dr  J.  L.  Hickman,  8-9  Nov  1955  - 
TM  :  K300-30I.  Mt  Wellington  (type-locality)  I47°I5'E.  42°55'S.  Map,  14/4  :  under  stones, 
creek,  Bett's  Vale,  Dr  J.  L.  Hickman,  4  Mar  1954  -  ™  :  K3°2  >  Shoobridge  Bend  Track, 
approx.  580  m  altitude,  in  loam  and  clay  in  Eucalypt-fern  woodland,  B.  G.  M.  Jamieson  and 
E.  Bradbury,  19  Aug  1971  -TM  :  1X303.  Lenah  Valley,  near  Newton  Falls,  i47°2o'E.  42°5o'S. 
Map,  14/5,  Dr  J.  L.  Hickman,  24  Jun  1957  -  TM  :  1X304.  Mt  Nelson,  Sandy  Bay,  i47°2o'E. 
42°55'S.  Dr  J.  L.  Hickman,  n  Sep  1953  -  TM  :  1^305.  Risdon,  i47°2o'E.  42°5o'S.  Map, 
14/6,  Professor  V.  V.  Hickman,  26  Jun  1947  -  TM  :  1^306-307  ;  East  Risdon,  from  under 
stones  on  hill  and  in  valley,  Dr  J.  L.  Hickman,  14  Aug  1954  -  BM(NH)  :  1972.8.15-17.  Eagle- 
hawk  Neck,  under  bark,  i47°55'E.  43°oo'S.  Map,  19/2,  Dr  J.  L.  Hickman  1954  -  AM  :  W5I94- 
5198  ;  BJ  :  Ti6-i8.  Tinderbox,  under  fallen  eucalyptus  leaves,  i47°2o'E.  43°O5'S.  Map, 
19/3,  Dr  J.  L.  Hickman,  4  Aug  1957  -  TM  :  KsoS. 

REMARKS.  The  above  account  of  the  lectotype  confirms  and  considerably  aug- 
ments Spencer's  description.  As  shown  in  the  table  of  field-variation,  the  accessory 


244 


B.   G.   M.    JAMIESON 


PQ 


* 

o 


O 


1 
S 


N   0,' 

IH          ' 

6 


TASMANIAN  EARTHWORMS  245 

genital  markings  were  more  numerous  in  the  material  described  by  Spencer  but  the 
evidence  offered  by  Jensz  and  Smith  (1969)  for  regarding  the  lectotype  which  they 
selected  as  part  of  the  type-series  is  satisfactory.  Location  of  paired  genital  markings 
in  intersegmental  furrow  16/17  is  almost  constant  for  the  species,  the  sole  exceptions 
in  the  material  examined  being  those  from  Goulds  County  (10/1).  The  latter  speci- 
mens are  also  exceptional  in  having  genital  markings  in  20/21  (seen,  however,  in  the 
right  side  in  a  Sandy  Bay  specimen  (14/5))  and  have  a  very  large  gizzard.  The 
possibility  that  these,  and  perhaps  specimens  from  other  localities,  are  subspecifically 
or  specifically  distinct  from  the  type-population  deserves  further  investigation  but 
on  present  evidence  recognition  of  distinct  taxa  is  not  justified. 

Perionychella  (Vesiculodrilus)  lacustris  (Stephenson,  1924) 

Fig.  i,  76,  16! 

Perionyx  lacustris  Stephenson,  1924  :  546-547. 
Diporochaeta  lacustris  :   Jamieson,  I97ib  :  83. 

1  =  45-60  mm,  w  =  2-4-3-0  mm,  s  =  95-107. 

Prostomium  epilobous  f,  wedge-shaped  (type)  or  tanylobous  (Bronte).  First 
dorsal  pore  4/5.  Setae  on  XII  18  (type),  22  (Stephenson),  14  and  16  (Bronte)  ; 
caudally  (Bronte)  16  and  23  ;  20  on  V  and  XIX,  22  on  IX  (Stephenson).  The  dorsal 
setal  gap  regular  anteriorly  and  in  the  midbody  equalling  2-$yz,  but  irregular  and 
indistinct  caudally  ;  ventral  gap  regular  throughout,  equalling  or  >  2ab  anteriorly. 
Clitellum  annular  |XIII-|XVII.  Male  pores  in  b  on  prominent  ovoid  papillae 
occupying  ac  on  XVIII.  Paired  eye-like  markings  in  be  in  16/17,  I7/1^  and  18/19, 
those  of  17/18  slightly  median  of  the  others ;  a  median  sucker-like  pad  postsetal 
ventrally  in  each  of  VII  and  VIII  ;  an  indistinct  glandular  tumescence  present  on 
each  side  in  ac  on  XVII  and  XIX.  Female  pores  paired  in  XIV  anteromedian  of  a. 
Spermathecal  pores  3  pairs,  in  6/7  to  8/9,  on  small  papillae  in  cd,  closer  to  c. 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  in  VIII-XII,  well 
developed.  Gizzard  small,  almost  vestigeal,  in  V,  hidden  in  septal  glands.  Calci- 
ferous  glands  absent  but  in  the  type  the  oesophagus  is  swollen  and  vascularized,  and 
has  low  internal  rugae,  in  XIV  and  XV.  Intestinal  origin  |XVII ;  typhlosole  absent. 
Nephridia  simple,  stomate,  vesiculate,  exonephric  holonephridia  commencing  (but 
rudimentary  ?)  in  II  (preseptal  funnel  demonstrated  in  XIV)  ;  bladders  large,  sub- 
spherical,  tapering  ectally  or,  in  the  type,  represented  at  most  by  slight  intraparietal 
dilatations  of  the  terminal  duct.  Holandric  (funnels  iridescent)  ;  gymnorchous  ; 
seminal  vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  large  in  XIV. 
Prostates  tubuloracemose,  in  XVII  to  XX  (type)  or  to  XXI,  XXII  (Bronte)  ;  only 
partially  resolvable  into  a  simple  linear  arrangement ;  depressed,  deeply  incised  and 
tabulated ;  vas  deferens  (see  in  Bronte  specimens  only)  joining  gland  near  its 
muscular  duct.  Penial  setae  delicate,  filiform,  sharply  pointed  ;  apical  extremity 
rounded  ;  ornamentation  absent ;  length  ?  Penial  setae  present  in  the  type  but  not 
examined.  Spermathecae  3  pairs ;  diverticulum  (inseminated)  single,  clavate, 
uniloculate. 


246  B.   G.  M.   JAMIESON 

TYPE-LOCALITY.     Great  Lake,  under  stones  in  water  at  margin  of  the  lake, 
i46°45'E.  4i°55'S.    Map,  8/2. 
MATERIAL  EXAMINED. 

Syntypes,  BM(NH)  :  1924.10.21.1-5,  of  which  one,  referred  to  as  'type'  in  the  above  account, 
was  closely  examined. 

Lyell  Highway,  5  miles  from  Bronte  towards  Hobart,  i46°35'E.  42°i5'S.  Map,  13/7,  Dr  J.  L. 
Hickman,  24  May  1954  ~  tw°  specimens  TM  :  K265  and  BM(NH)  :  1972.8.2. 

REMARKS.  The  new  description  of  a  type-specimen  in  the  above  account  con- 
siderably augments  Stephenson's  description.  In  this  type-specimen,  which  had  not 
been  previously  dissected,  and  in  the  single  one  of  the  type-specimens  which  had 
previously  been  dissected,  the  following  observations  of  Stephenson  are  not  confirmed 
and  are  considered  incorrect :  clitellum  in  XIV-XVI ;  gizzard  in  VI ;  intestinal 
origin  (about)  XVIII ;  penial  setae  absent.  Genital  markings  are  similar  in  the 
two  specimens  and  include  midventral  markings  in  VII  and  VIII. 

P.  (V.)  montisarthuri  is  morphologically  close  to  this  species  but  shows  sufficient 
differences  to  be  considered  a  distinct  species.  These  differences  include  the  setal 
counts,  the  bilobed  nephridial  vesicles,  the  sessile  spermathecae  and  the  precise 
configuration  of  the  genital  field. 

Perionychella  (Vesiculodrilus)  montisarthuri  sp.  n. 

Fig.  i,  I2A,  15!!,  168.     Table  2 
1  =  79  mm,  w  =  3-6  mm,  s  =  86. 

Prostomium  epilobous  \,  open.  First  dorsal  pore  3/4.  Setae  on  XII  8  (10  on  XI)  ; 
caudally,  14  or  16  per  segment  "  the  dorsal  gap  regular  anteriorly  and,  in  the  mid- 
body,  equalling  2,yz,  irregular  caudaUy  ;  ventral  gap  regular  throughout,  equalling 
2ab  anteriorly.  Nephropores  in  a  single  straight  series  on  each  side  in  setal  line  3 
anteriorly  and  in  the  mid-body ;  caudally  in  s.l.  4-5.  Clitellum,  annular  £XIII- 
XVI.  Male  pores  in  b  on  small,  flat,  ovoid  markings  on  XVIII ;  two  ovoid  genital 
markings  occupy  the  space  from  a  to  a  little  lateral  of  b  at  17/18  and  18/19.  Female 
pores  paired  on  XIV,  anterior  and  slightly  median  to  a.  Spermathecal  pores  3  pairs 
in  b  in  6/7-8/9,  on  small  papillae  each  of  which  is  succeeded  by  a  small  ovoid  marking 
on  the  posterior  segment. 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  in  VIII-XII,  well 
developed.  Gizzard  in  V,  moderately  large  but  only  weakly  muscular.  Calciferous 
glands  absent.  Oesophagus  with  circumferential  vascular  striae  in  VII-XIV, 
widest  in  XIV  and  XV  in  which  internally  there  are  numerous  radial  lamellae  which 
almost  occlude  the  lumen  ;  narrow  in  XVI  in  which  there  are  a  few,  low  radial  folds. 
Intestinal  origin  |XVII  ;  typhlosole  absent.  Nephridia  simple  stomate,  vesiculate, 
exonephric  holonephridia  commencing  in  II  (funnels  demonstrated  in  the  forebody)  ; 
bladders  at  first  elongate  avoid  but  by  VI  with  a  suggestion  of  a  lateral  basal  lobe  ; 
thereafter  the  lateral  lobe  increases  in  size  relative  to  the  median  portion  (the  latter 
receiving  the  nephridial  duct)  until,  by  about  XIX,  the  bladder  is  broader  than  long 
and  consists  of  two  approximately  equal  lobes,  the  lateral  lobe  constituting  a  broadly 


TASMANIAN  EARTHWORMS  247 

sessile  diverticulum  ;  caudally  the  bladders  are  less  distinctly  bilobed.  Holandric 
(funnels  iridescent  in  X  and  XI),  gymnorchnous  ;  seminal  vesicles  racemose,  in  IX 
and  XII.  Metagynous  ;  ovisacs  small  in  XIV.  Prostates  depressed  very  tortuous 
tubes,  in  XVII-XXI ;  double  vas  deferens  joining  the  gland  shortly  ental  of  the 
slender,  sinuous  muscular  duct.  Penial  setae  (holotype)  slender,  strongly  curved, 
almost  filiform  with  delicate  thinner,  elongate  tip  ;  ensheathed  in  setal  follicle  and 
not  suitable  for  electron  microscopy ;  length  (?),  incomplete.  Spermathecae  3 
pairs,  diverticulum  (inseminated)  single,  clavate,  uniloculate,  exceptional  in  joining 
the  ectal  end  of  the  broad  subspherical  ampulla ;  ampulla  almost  sessile  on  the 
parietes. 
MATERIAL  EXAMINED. 

Mt  Arthur,  from  rainforest,  i47°2o'E.  4i°i5'S.  Map,  9/1,  Mr  A.  J.  Dartnall  and  Mr  R.  C. 
Kershaw,  15  Oct  1971  -  holotype  TM  :  Ksog. 

REMARKS.    This  species  is  morphologically  close  to  P.  (V.)  hellyeri  (q.v.) 

Perionychella  (Vesiculodrilus)  mortoni  (Spencer,  1895) 
Fig.  i,  I2B,  I5E-G,  i6T  ;  Plates  99-102.     Table  2 

Cryptodrilus  mortoni  Spencer,  1895  :  36-37,  PI.  i,  fig.  7-9. 

Plutellus  mortoni  ;  Michaelsen,  1900  :  176. 

Woodwardiella  mortoni ;   Jamieson,  1970  :  104,  105  ;   19710  :  92. 

Dimensions  (see  Variation).  Anatomy  (specimen  i,  locality  14/4,  19  Aug  1971)  : 
Prostomium  canaliculate,  tanylobous.  First  dorsal  pore  4/5.  Setae  in  8  regular 
longitudinal  rows  throughout.  Nephropores  conspicuous,  commencing  anteriorly  in 
II  in  c  lines  but  shortly  below  c  lines  behind  the  midclitellum.  Clitellum  annular, 
XIII-XVII,  strongly  developed  and  pigmented  in  XIV-XVII,  unpigmented  and 
weakly  developed  in  XIII ;  possibly  some  slight  clitellar  modification  in  XVIII. 
Male  pores  on  small  papillae  in  ab  of  XVIII.  Accessory  genital  markings  :  paired 
elliptical  pads  with  depressed  centres  in  ab  in  17/18,  19/20,  20/21  and  21/22,  those  in 
17/18  with  centres  slightly  lateral  of  those  of  the  other  markings  and  truly  inter- 
segmental ;  the  markings  in  19/20-21/22,  though  intersegmental  actually  postsetal 
in  XIX-XXI,  respectively ;  a  median  elliptical  pad  present  in  17/18  bridging  the 
paired  markings  of  the  intersegment.  Median  markings  each  with  the  form  of  a 
depressed  transverse  intersegmental  band  with  anterior  rim  in  aa  in  5/6-8/9.  Female 
pores  anteromedian  of  setae  a  of  XIV,  midway  between  the  setal  arc  and  anterior 
border  of  the  segment  in  a  common  oval  field.  Spermathecal  pores  5  pairs  on 
minute  circular  papillae  immediately  in  front  of  intersegmental  furrows  4/5-8/9,  in 
a  lines. 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  well  developed,  in 
VIII-|XIII.  Gizzard  very  large  and  firm,  in  V  but  projecting  posteriorly  to  the 
level  of  XL  Calciferous  glands  absent.  Oesophagus  strongly  vascularized  and 
increasingly  dilated  in  XIII-XVI  with  well-developed  internal  vascular  rugae, 
especially  in  XV  and  XVI.  Intestinal  origin  XVIII ;  typhlosole  absent.  Neph- 
ridia  stomate,  vesiculate  exonephric  holonephridia  commencing  in  II  (preseptal 


248  B.   G.   M.   JAMIESON 

funnel  demonstrated  for  those  of  II)  ;  tortuously  coiled  but  simple,  none  tufted  ; 
bladders  at  first  elongate,  wide  tubes,  each  of  slightly  irregular  diameter ;  by  XII 
bent  midway  at  a  right  angle  ;  thereafter  with  a  lateral  diverticulum  extending  from 
the  bend,  the  portion  ectal  to  the  diverticulum  being  eliminated  after  a  few  segments. 
Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles  race- 
mose in  IX  and  XII.  Metagynous ;  ovisacs  present.  Prostates  much  coiled 
depressed  tubes,  in  XVIII-XXI ;  vas  deferens  joining  the  duct  shortly  ectal  of  the 
gland.  Penial  setae  short  and  moderately  stout,  the  tip  irregular,  widened,  roughly 
spatulate  and  bent  a  little  or  through  a  right  angle,  this  apical  modification  visible 
under  the  light  microscope  ;  shaft  ornamented  ectally  with  a  few  groups  of  anteriorly 
directed  scarcely  protuberant  teeth  regularly  spaced  along  it ;  the  surface  of  the  seta 
undercut  beneath  them  ;  length  0-5  mm  (3  setae,  Mt  Wellington  and  East  Risdon 
specimens,  as  illustrated).  Spermathecae  5  pairs ;  diverticulum  (inseminated) 
single,  clavate,  uniloculate. 

Variation.  A  minimum  of  two  specimens  from  each  of  localities  7/2,  9/2,  14/3, 
14/4,  14/5,  14/6  and  the  single  specimen  from  19/2  were  examined  for  variation  in 
external  characters.  Variation  in  internal  anatomy  is  described  for  at  least  one 
specimen  from  each  locality. 

Genital  markings.  Postsetal  genital  markings  were  present  in  aa  in  each  of 
segments  IV- VIII  in  all  specimens  but  in  those  from  Fern  Glade  (locality  7/2)  these 
were  exceptional  in  being  paired.  Paired  markings  were  present  in  17/18  and  post- 
setally  in  each  of  segments  XIX,  XX  and  XXI,  with  the  exception  that  those  in  XXI 
were  absent  in  one  of  the  two  specimens  from  each  of  localities  9/2,  14/3  and  14/5 
and  from  most  East  Risdon  specimens  (14/6).  The  median  genital  marking  in  17/18, 
though  characteristic  of  the  species,  is  absent  from  most  but  not  all  of  the  East 
Risdon  specimens. 

Spencer  did  not  observe  genital  markings  in  IV- IX  (these  are  often  faint  in  the  new 
material)  but  he  depicted  median  and  paired  markings  in  17/18,  as  here  described, 
and  paired  markings  in  intersegments  18/19  and  19/20.  The  latter  two  pairs  were 
presumably  postsetal  rather  than  intersegmental  and  it  appears  probable  from  the 
present  study  that  they  lay  in  segments  XIX  and  XX. 

Other  variation.  1  =  56-212  mm,  w  =  3-5-6-5  mm,  s  =  113-274.  Prostomium 
epilobous  |-f,  epitanylobous  or  tanylobous  ;  often  canaliculate.  First  dorsal  pore 
usually  in  4/5,  rarely  in  3/4  or  5/6.  Body  strongly  canaliculate  to  not  canaliculate. 

Variation  from  the  description  in  internal  characters  occurs  in  the  intestinal  origin 
which  is  XVII  in  9/2  and  19/2  ;  in  the  presence  of  a  low  dorsal  intestinal  ridge, 
though  no  true  typhlosole  is  present,  in  9/2  and  14/6  ;  and  in  the  absence  of  diverti- 
cula  of  the  nephridial  bladders  in  9/2.  Absence  of  nephridial  diverticula  is  especially 
noteworthy  and  is  a  variation  hitherto  unknown  intraspecifically  in  otherwise  diverti- 
culate  species.  At  present  there  is  insufficient  justification  for  taxonomic  recognition 
of  the  variant. 

MATERIAL  EXAMINED. 

Approximately  100  sexual  specimens  were  examined  from  the  following  localities  :  Fern 
Glade,  Emu  River,  Burnie,  I45°55'E.  4i°o5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  24  Aug  1954. 
Mt  Arthur  (east),  wet  sclerophyll  forest,  i47°2o'E.  4i°i5'S.  Map,  9/1,  Mr  A.  J.  Dartnall  and 


TASMANIAN  EARTHWORMS 


249 


.m 


1mm 


sp.p  3 


sp.p  5 


1mm 


FIG.  13.     Genital  fields. 


B 

Perionychella  (Vesiculodrilus)  obliquae  :  A,  holotype  ;  B,  17/1, 
AM  :  W52O2. 


Mr  R.  C.  Kershaw,  15  Nov  1971.  St  Columba  Falls,  I47°55'E.  4i°2o'S.  Map,  9/2,  Dr  J.  L. 
Hickman,  17  Apr  1954.  Collinsvale,  near  Hobart,  in  Myrtle  Forest,  i47°io'E.  42°5o'S.  Map, 
14/3,  Dr  J.  L.  Hickman,  8-9  Nov  1955.  Mt  Wellington,  I47°I5'E.  42°55'S.  Map,  14/4, 
Dr  J.  L.  Hickman,  13  Jan  1954  •  Dr  J-  L.  Hickman,  27  Oct  1955  ;  Mr  W.  Radford,  Apr  1952  ; 
Professor  V.  V.  Hickman,  28  Jun  1947  ;  13  Sep  1951  -  B  J  :  Tig-20  ;  Betts  Vale,  Dr  J.  L. 
Hickman,  4  Mar  1954  -AM  '•  W5 199-5201  ;  Shoobridge  Bend  track,  approx.  580  m,  Dr  B.  G. 
M.  Jamieson  and  Mr  E.  A.  Bradbury,  19  Aug  1971  -  BM(NH)  :  1972.8.18-25.  Hobart,  Water- 
works Road,  under  stones,  i47°2o'E.  42°5o'S.  Map,  14/5,  Dr  J.  L.  Hickman,  17  Aug  1954. 
Lenah  Valley,  track  along  Newtown  Creek,  Dr  J.  L.  Hickman,  15  Sep  1953  ;  Lenah  Valley, 
under  stones  near  Newtown  Falls,  Dr  J.  L.  Hickman,  24  Jun  1957  -  BM(NH)  :  1972.8.26-30; 

15 


250  B.   G.   M.   JAMIESON 

Sandy  Bay,  Dr  J.  L.  Hickman,  Sep  1954  ;  Domain,  14  Aug  1954.  East  Risdon,  under  stones 
on  hill  and  in  valley,  i47°2o'E.  42°5o'S.  Map,  14/6,  Dr  J.  L.  Hickman,  14  Aug  1954  -  BJ  :  T2I  - 
22  ;  Risdon,  Professor  V.  V.  Hickman,  26  Jun  1947.  Eagle  Hawk  Neck,  i47°55'E.  43°oo'S. 
Map,  19/2,  Dr  J.  L.  Hickman,  13  May  1954.  Tasmania  :  T.M.  I5527/K97,  Dr  J.  L.  Hickman. 
All  specimens  lodged  in  the  Tasmanian  Museum  (K97,  K4O2-4I9)  except  where  indicated  above. 

REMARKS.  This  species  and  P.  (V.)  hobartensis  are  the  most  widespread  and 
commonly  found  megascolecid  earthworms  in  Tasmania  and  are  morphologically 
close.  A  notable  difference  is  location  of  the  paired  genital  markings  at  the  posterior 
borders  of  their  segments  in  mortoni  whereas  in  hobartensis  they  are  intersegmental. 

Perionychella  (Vesiculodrilus)  obliquae  sp.  n. 
Fig.  i,  I3A,  B,  i6U,  V 

1  =  115  mm  (holotype  ;  paratype  i  is  a  posterior  amputee),  w  =  6-6,  6-0  mm, 
s  =  118. 

Prostomium  epilobous  £,  open.  First  dorsal  pore  3/4.  Setae  on  XII  27,  25  ; 
caudally  40,  38  ;  ventral  break  distinct  throughout,  equal  to  2ab  in  forebody  and  $ab 
posteriorly ;  dorsal  gap  not  distinct  caudally  being  equal  to  dorsal  couple  (yz)  ; 
distinct  anteriorly,  equalling  2yz.  Nephropores  difficult  to  observe,  anteriorly  in 
setal  lines  6  and  posteriorly  in  s.l.  8  (holotype)  or  8-9  (paratype  i).  Clitellum 
annular,  |XII-XVII.  Male  pores  on  small  papillae  in  b  lines.  Five  pairs  of  eye-like 
markings  on  the  anterior  margins  of  segments  XX  and  XXI  in  b  lines  and  on  segments 
XVII,  XVIII  and  XIX  in  ab.  Female  pores  paired  anterior  and  a  little  median  to 
a  on  XIV.  Spermathecal  pores  5  pairs,  in  4/5-8/9  in  ab  on  small  papillae. 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  in  VII-XIII,  very 
well  developed.  Gizzard  small,  almost  vestigial,  in  V.  Extramural  calciferous 
glands  absent ;  intestinal  origin  |XVII ;  typhlosole  absent.  Nephridia  simple 
exonephric,  vesiculate  holonephridia  commencing  in  II ;  preseptal  funnels  present 
at  least  from  III  posteriorly  ;  bladders  long,  tubular  or  somewhat  dilated,  bent  into 
a  U  or  V  shape.  Holandric  (funnels  iridescent)  ;  gymnorchous  ;  seminal  vesicles 
racemose,  in  IX  and  XII.  Metagynous  ;  large  ovisacs  in  XIV.  Prostates  depressed 
tubular,  somewhat  lobulated,  with  several  bends  compacted  into  a  circular  outline 
and  spuriously  appearing  racemose,  in  XVII-XIX.  A  small  follicle  but  no  seta 
present  in  a  of  XVIII  in  the  holotype  ;  penial  seta  from  AM  :  W5202  very  slender, 
straight,  unornamented  (under  light  microscope),  length  =  0-58  mm  ;  a  vestigial 
seta  from  paratype  under  the  electron  microscope  displays  no  ornamentation  ;  tip 
pointed,  aquiline,  length  =  0-34  mm.  Spermathecae  5  pairs  ;  diverticulum  (in- 
seminated) single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

Obliqua-iorest,  2  miles  inland,  south  of  Interview  River,  I44°55'E.  4i°35'S.  Map,  6/1,  Mr 
W.  Jackson,  31  Dec  1953  -  holotype  TM  :  KSIO  ;  paratype  i  BM(NH)  :  1972.8.31.  Port 
Davey,  Kelly's  Beach,  I45°55'E.  43°2o'S.  Map,  17/1,  Tasmanian  Biological  Survey  :  J 17, 
Jan  1940  -  AM  :  W5202. 

REMARKS.  Differences  from  the  Interview  River  type-specimens  shown  by  the 
Port  Davey  specimen  are  as  follows  :  clitellar  width  2-7  mm  ;  first  dorsal  pore  4/5  ; 


TASMANIAN  EARTHWORMS  251 

setae  22  in  segment  XII ;  accessory  genital  markings  absent  at  17/18  and  18/19  » 
a  midventral  circular  genital  marking  present  posteriorly  in  VII  (possibly  an  arte- 
fact) ;  gizzard  moderate  in  size  ;  intestinal  origin  XVII,  not  |XVII ;  nephridial 
bladders  subspherical  though  ectally  tapering.  These  differences  do  not  appear 
sufficient  for  recognition  of  a  distinct  species  for  the  Port  Davey  material,  especially 
as  only  a  single  specimen  is  available.  Larger  series  from  both  localities  are  needed. 
The  genital  field  of  this  species  is  reminiscent,  though  substantially  different  from, 
that  of  P.  (V.)  hobartensis  which  obliquae  closely  resembles.  It  differs  conspicuously 
from  hobartensis  and  from  P.  (V.)  tunnackensis,  to  which  it  is  also  morphologically 
close,  in  the  perichaetine  arrangement  of  its  setae. 


Perionychella  (Vesiculodrilus)  pedderensis  sp.  n. 

Fig.  i,  I2C,  i6W 
1  =  50  mm,  w  =  1-6  mm,  s  =  123. 

Prostomium  canaliculate,  indistinctly  epilobous  £,  open.  Dorsal  pores  absent. 
Forebody  dorsoventrally  depressed  ;  hindbody  with  wide  dorsal  gutter ;  anus 
terminal  but  deeply  incising  the  pygidium  dorsally  and  ventrally.  Setae  10  per 
segment  anteriorly,  increasing  to  12  per  segment  caudally,  rows  regular  ;  dorsal  and 
ventral  gaps  large,  the  dorsal  larger.  Nephropores  ?  Clitellum  annular,  XIII-XVI. 
Male  pores  in  ab  of  XVIII  on  small  papillae.  Accessory  genital  markings  :  a  mid- 
ventral  tumid  pad  traversing  bb  in  each  of  intersegments  17/18  and  18/19,  eacn  Pad 
with  a  deep  groove  corresponding  with  the  intersegment ;  paired  eye-like  genital 
markings  centred  in  a  lines  in  19/20  and  20/21  (that  on  the  left  in  20/21  barely  per- 
ceptible) and  posteriorly  in  VIII  shortly  lateral  of  a  lines.  Female  pores  at  |  aa, 
midway  between  the  setal  arc  and  the  anterior  border  of  XIV.  Spermathecal  pores 
4  pairs,  minute,  in  5/6  shortly  lateral  of  a,  in  6/7  in  ab,  in  7/8  and  8/9  in  b. 

Last  hearts  XII.  Suboesophageal  VII I -XIII.  Gizzard  moderate,  in  VI. 
Oesophagus  simple  ;  intestinal  origin  XVII  ;  typhylosole  absent.  Holonephridia 
commencing  in  II  but  appearing  rudimentary  to  VII ;  each  with  a  thick-walled 
subspherical  bladder  discharging  presetally  in  c  line  ;  preseptal  funnel  well  developed. 
Holandric  (funnels  iridescent)  ;  gymnorchous  ;  seminal  vesicles  racemose,  in  IX 
and  XII.  Metagynous  ;  large  ovisacs  in  XIV.  Prostates  thickly  tubular,  i  pair, 
in  XVIII-XIX,  or  XX  ;  vas  deferens  joining  ectal  limit  of  gland.  Penial  setae 
present  (holotype)  but  structure  not  elucidated  ;  no  visible  ornamentation  ;  length 
(incomplete  ?)  =  0-41  mm.  Spermathecae  4  pairs  ;  diverticulum  (inseminated) 
single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

Lake  Pedder,  sorted  from  interstitial  fauna,  i46°io'E.  42°55'S.  Map,  13/2,  Mr  D.  Tyler, 
25  Feb  1971  -holotype  TM  :  K$II. 

REMARKS.  The  absence  of  dorsal  pores,  which  is  unique  in  the  genus  in  Tasmania, 
and  the  reduction  of  the  anterior  nephridia  are  elsewhere  correlated  with  an  aquatic 
existence  in  megadriles. 


252 


B.   G.   M.   JAMIESON 


XII 


XVII 


VIII 


IX 


1mm 


sp.p 


B 


1mm 


FIG.  14.     Genital  fields.    Perionychella  ( Vesi culodrilus)  t unnackensis,  holotype  :   A, 
male  field  ;   B,  spermathecal  field. 


TASMANIAN  EARTHWORMS  253 

Perionychella  (Vesiculodrilus)  tunnackensis  sp.  n. 

Fig.  i,  I4A,  B,  i6X.     Table  2 
1  =  33.  35  mm>  w  =  1-4,  1-3  mm,  s  =  82,  81. 

Prostomium  faintly  canaliculate,  epilobous  £.  First  dorsal  pore  4/5.  Setae  8 
per  segment  in  regular  longitudinal  rows  throughout.  Nephropores  in  c.  Clitellum 
annular,  13-16.  Male  pores  in  ab  on  small  papillae  on  18,  the  two  papillae  joined  by 
a  low  median  ridge,  intersegmental  ridges  at  17/18  and  18/19  flank  the  male 
papillae  and  correspond  in  extent  to  the  ridge  between  the  papillae.  Paired  eye-like 
transversely  conjoined  genital  markings  in  ab  at  19/20  (holotype,  paratype)  and  20/21 
(holotype)  ;  post-setal  eye-like  markings  occur  with  centres  in  a  in  VII  (paired  in 
paratype  ;  left  only  in  holotype)  ;  VIII  (paired)  and  IX  (paired  in  paratype  I  ; 
right  only  in  holotype).  Female  pores  a  pair  anteromedian  of  a  on  XIV.  Sperma- 
thecal  pores  3  pairs  in  a  at  6/7,  7/8  and  8/9. 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  not  distinguishable 
from  the  well-developed  oesophageal  plexus.  Gizzard  in  V,  moderate  to  small, 
muscular  but  easily  compressible.  Calciferous  glands  absent.  Oesophagus  with 
circumferential  vascular  striae  in  (VII  ?),  VIII-XV  ;  in  X-XV  with  at  first  few  and 
small,  posteriorly  increasingly  numerous  and  larger  longitudinal  rugae  which  in  XIV 
and  XV  approach  the  appearance  of  laminae.  Intestinal  origin  XVII ;  typhlosole 
absent.  Nephridia  simple  stomate,  vesiculate  holonephridia,  commencing  in  II 
(funnels  traced  at  least  from  IV)  ;  bladders  subspherical,  tapering  to  the  pore  ;  large 
in  all  but  a  few  anteriormost  segments.  Holandric  (funnels  iridescent  in  X  and  XI)  ; 
gymnorchous  ;  seminal  vesicles  racemose  in  IX  (?)  and  XII.  Metagynous  ;  ovisacs 
large,  in  XIV.  Prostates  depressed  tubular  with  incised  adpressed  coils,  in  XVIII- 
XX  ;  vas  deferens  joining  the  gland  shortly  ental  to  the  muscular  duct.  Penial 
setae  (holotype)  filiform  ;  tip  pointed  and  slightly  upturned  ;  no  ornamentation  ; 
length  =  0-76  mm.  Spermathecae  3  pairs,  decreasing  in  size  anteriorly ;  diverti- 
culum  (uninseminated)  single,  clavate  uniloculate. 

MATERIAL  EXAMINED. 

Tunnack,  under  logs  and  stones,  damp  conditions,  i47°3o'E.  42°25'S.  Map,  14/2,  Dr  J.  L. 
Hickman,  18  Aug  1954  -  holotype  TM  :  K3I2  ;  paratype  BM(NH)  :  1972.8.32. 

REMARKS.  As  in  all  Tasmanian  species  of  the  genus,  the  genital  markings  of  this 
species  are  distinctive.  It  is  closest  to  P.  (V.)  hobartensis  from  which  the  3  pairs  of 
spermathecal  pores  are  an  additional  distinction. 


Perionychella  (subgenus  ?)  bassana  (Spencer,  1895) 

Megascolides  bassanus  Spencer,  1895  :  46-47,  PI.  Ill,  fig.  34-36. 
Plutellus  bassanus  ;  Michaelsen,  1900  :  169  ;   Jamieson,  ig7ic  :  87. 

1  =  88  mm,  w  >  3  mm,  s  ?. 

Prostomium  prolobous.     First   dorsal  pore  in  4/5  (?).     Setae  8   per  segment. 
Nephropores  in  c  lines.     Clitellum  saddle-shaped,  embracing  XIV-XIX,  ventral 


254 


B.   G.   M.   JAMIESON 


FIG.  15.  Nephridial  bladders  in  Perionychella.  A  &  B,  Perionychella  (Vesiculodrilus) 
hobartensis  :  A,  anterior  segments  ;  B,  an  intestinal  segment.  C,  P.  (V.)  evansi,  holoytpe, 
D,  P.  (V.)  glandifera,  holotype.  E-G,  P.  (V.)  mortoni,  14/4,  BM(NH)  :  1972.8.18. 
H,  P.  (V.)  montisarthuri,  holotype,  in  situ,  showing  setae  c.  (A  &  B,  left  bladders  ;  others, 
right  bladders.) 


margins  in  ab  excepting  the  anterior  part  of  XIV,  the  whole  of  which  is  included. 
Male  pores  on  papillae  in  XVIII  in  a  lines.  Accessory  genital  markings  :  a  median 
ventral  patch  on  each  of  XVII  and  XVIII ;  and  2  papillae  in  XIX  in  a  lines,  con- 
joined by  a  median  transverse  ridge.  Female  pores  on  XIV  within  the  tumid  annular 
part  of  the  clitellum  near  the  midventral  line.  Spermathecal  pores  2  pairs,  in  7/8 
and  8/9,  in  ab. 

Dorsal  blood  vessel  single  ;  last  hearts  in  XIII ;  no  continuous  supra-oesophageal. 
Gizzard  in  V.  True  calciferous  glands  absent  but  vascular  swellings  in  XIII  and 
XIV  ;  intestinal  origin  in  XIX.  Nephridia  stomate  holonephridia.  Holandric  ?, 
sperm  funnels  in  X  and  XL  Seminal  vesicles  on  anterior  walls  of  X  (?),  XI  and  XII. 
Metagynous  ;  ovisacs  ?.  Prostates  tubular,  small  and  coiled,  in  XVIII.  Sperma- 
thecae  2  pairs,  in  VIII  and  IX  ;  the  diverticulum  simple  and  less  than  half  the  length 
of  the  sac. 

TYPE-LOCALITY.     King  Island  in  Bass  Strait. 


TASMANIAN  EARTHWORMS  255 

MATERIAL  EXAMINED. 

None.     The  types  are  missing  from  the  National  Museum  of  Victoria. 

REMARKS.  It  is  not  entirely  certain  that  this  species  should  be  assigned  to 
Perionychella.  The  saddle-shaped  clitellum  is  exceptional,  and  presence  or  absence 
of  a  typhlosole  is  not  recorded,  but  otherwise  nothing  in  the  brief  description  ex- 
cludes it  from  the  genus. 


Perionychella  (subgenus  ?)  decatheca  (Michaelsen,  1910) 
Plutellus  decatheca  Michaelsen,  1910  :  81-83,  Fig-  XIV-XVI ;   Jamieson,  19710  :  87. 
1  =  100  mm,  w  =  10  mm,  s  =  164. 

Prostomium  epilobous  \,  closed.  First  dorsal  pore  in  4/5.  Setae  widely  paired  ; 
in  the  forebody  aa  :  ab  :  be  :  cd  :  dd  =  1-5:1:2:1-5:  4-5  ;  dd  =  o-6w  ;  in  the 
hindbody  1-5:1:1-3:1:  2-5  ;  dd  =  o-25«.  Clitellum  annular,  XIV-XVIII.  Male 
pores  on  small  papillae  in  XVIII,  approximately  in  ab.  Accessory  genital  markings  : 
paired  transverse  almost  linear  papillae  anterior  and  posterior  to  the  male  papillae, 
in  XVII,  XIX,  XX  and  XXI  extending  medianly  over  a  lines  and  laterally  over  b 
lines  ;  the  markings  on  XVII  much  larger  than  those  of  XIX-XXI,  almost  contigu- 
ous medianly  and  nearly  obscuring  the  male  papillae.  Female  pores  indicated  by  a 
long  transverse  furrow  anteriorly  in  XIV.  Spermathecal  pores  5  pairs,  in  4/5-8/9, 
in  a  lines. 

Dorsal  blood  vessel  single  ;  last  hearts  in  XIII.  A  large  gizzard  in  V.  Calciferous 
glands  absent ;  oesophagus  in  XIV  and  XV  appearing  more  vascular  than  elsewhere, 
but  only  slightly  swollen.  Holonephric.  Holandric  ;  seminal  vesicles  racemose  in 
IX  and  XII.  Prostates  tubular  ;  duct  short  and  thin.  Penial  setae  ca.  1-2  mm 
long,  and  18  p,m  thick,  almost  straight,  proximally  slightly  curved,  distally  only 
slightly  tapered.  Distal  tip  rather  stout,  bent  into  a  small  hook.  Distal  end  of  the 
seta  ornamented  with  wide  deep  teeth,  which  project  very  steeply  from  the  seta ; 
the  teeth  lying  in  4  fairly  regular  spirally  arranged  lines.  Spermathecae  with  single, 
clavate,  uniloculate  diverticulum. 

TYPE-LOCALITY.    The  vicinity  of  Hobart. 
MATERIAL  EXAMINED. 

None. 

REMARKS.  Michaelsen  considered  P.  decatheca  to  be  close  to  P.  insularis  (Spencer, 
1895)  which  is  now  a  junior  synonym  of  P.  hobartensis.  Possibly  decatheca  is  merely 
a  variant  of  the  sympatric  hobartensis  but  differences  from  the  latter,  viz.  setal  rather 
than  intersetal  accessory  genital  markings  and  the  form  of  the  penial  setae  preclude 
union  of  the  two  taxa  on  present  information.  Location  of  the  genital  markings 
anteriorly  rather  than  posteriorly  in  their  segments  and  apparently  the  form  of  the 
penial  setae  exclude  decatheca  from  the  similarly  sympatric  and  not  dissimilar  P. 
mortoni.  Known  only  from  the  holotype. 


256 


B.   G.   M.   JAMIESON 


IffllR 


w 


FIG.  16.  Spermathecae  in  Perionychella.  A  &  B,  Perionychella  (P.)  capensis  :  A,  holotype 
(left  IX)  ;  B,  paratype  i  (left  IX).  C  &  D,  P.  (P).  hickmani  :  C,  holotype  (right  IX)  ; 
D,  paratype  3  (right  VIII).  E,  P.  (P.)  kershawi,  holotype  (right  VIII).  F,  P.  (P.)  weld- 
boroughi,  holotype  (right  IX).  G,  P.  (P.)  irregularis,  lectotype  (left  IX).  H,  P.  (7.) 
bithecata,  holotype  (left  IX).  I,  P.  (V.)  lacustris,  TM  :  K2&5  (left  IX).  J,  P.  (V.)  dil- 
wynnia,  13/4,  TM  :  Ka66  (right  IX).  K,  P.  (V.)  evansi,  holotype  (right  IX).  L  &  M, 
P.  (7.)  glandifera  :  L,  holotype  (left  IX)  ;  M,  paratype  (right  IX).  N  &  O,  P.  (V.) 
hellyeri :  N,  paratype  2  (right  IX)  ;  O,  holotype  (right  VIII).  P-R,  P.  (V.)  hobartensis  : 


TASMANIAN  EARTHWORMS  257 

Perionychella  (subgenus  ?)  ellisi  (Spencer,  1895) 
Fig.  i. 

Cryptodrilus  ellisii  (sic)  Spencer,  1895  :  42-43,  PL  2,  fig.  22-24. 
Plutellus  ellisi  ;  Michaelsen,  1900  :  172  ;   Jamieson,  19710  :  87. 

1  =  25-38  mm,  w  =  3  mm,  s  ?. 

Prostomium  epilobous  J,  it  and  the  body  canaliculate.  First  dorsal  pore  in  5/6. 
Setae  8  per  segment  throughout ;  be  =  cd  =  dd.  Clitellum  annular,  XIV-XVI  or 
posterior  XIII  also.  Male  pores  in  XVIII  in  b  or  perhaps  slightly  ventral  of  this,  on 
papillae.  Accessory  genital  markings  :  paired  elliptical  patches  in  ab  anteriorly  in 
X  and  XI ;  in  17/18  and  18/19  and  anteriorly  in  XX  and  XXI.  Female  pores 
anteromedian  of  setae  a  of  XIV.  Spermathecal  pores  3  pairs,  in  6/7-8/9,  in  b  lines. 
Last  hearts  in  XII.  Gizzard  in  V.  Sessile  paired  dorsolateral  calciferous  glands  in 
XIV  and  XV.  Intestinal  origin  in  XVII.  Holonephric.  Holandric ;  seminal 
vesicles  racemose,  in  IX  and  XII.  Prostates  tubular,  coiled,  in  XVII-XX.  Meta- 
gynous  ;  ovisacs  in  XIV.  Spermathecae  3  pairs  ;  diverticulum  short,  simple. 

TYPE-LOCALITY.     Dee  Bridge,  under  logs  and  stones,  I46°35'E.  42°i5'S.     Map, 
13/7- 
MATERIAL  EXAMINED. 

The  four  syntypes  (NMV  :  032)  were  found  to  be  in  poor  condition  and  no  longer  yield  useful 
information. 

Perionychella  (subgenus  ?)  moroea  (Spencer,  1895) 
Fig.  i. 

Perichaeta  moroea  Spencer,  1895  :  49,  PL  IV,  fig.  40-42. 
Diporochaeta  moroea  ;  Michaelsen,  1900  :  207  ;   Jamieson,  19710  :  83. 

1  =  zoo  mm,  w  ?,  s  ?. 

Prostomium  very  slightly  epilobous.  First  dorsal  pore  in  3/4.  Preclitellar  setae 
11-18  on  each  side  ;  intraclitellar  and  postclitellar  setae  21  per  side  ;  dorsal  and 
ventral  breaks  very  small.  Nephropores  in  setal  lines  9  or  10  in  the  midbody. 
Clitellum  ?  Male  pores  on  papillae  between  setal  lines  3  and  4.  Accessory  genital 
markings  :  median  ventral  ridge  on  XVIII  between  the  male  papillae  ;  a  pair  of 
elliptical  patches  in  ab,  in  19/20  (or  18/19  ?)•  Female  pores  on  XIV.  Spermathecal 
pores  2  pairs,  in  7/8  and  8/9,  in  ab. 

Last  hearts  in  XII ;  supra-oesophageal  present.  Gizzard  in  V.  True  calciferous 
glands  absent.  Intestinal  origin  XVII.  Holandric  ;  seminal  vesicles  racemose,  in 


P,  lectotype  (right  IX)  ;  Q,  14/1,  TM  :  K2g4  (right  IX)  ;  R,  14/6,  BM(NH)  :  1972.8.15 
(right  IX).  S,  P.  (V.)  montisarthuri,  holotype  (right  IX).  T,  P.  (V.)  mortoni,  14/4, 
BM(NH)  :  1972.8.18  (right  IX).  U  &  V,  P.  (V.)  obliquae  :  U,  holotype  (right  IX)  ;  V, 
17/1,  AM:  W5202  (right  IX).  W,  P.  (V.)  pedderensis,  holotype  (right  VIII).  X,  P. 
(V.)  tunnackensis,  holotype  (right  IX). 


258  B.   G.   M.   JAMIESON 

IX  and  XII.     Metagynous.     Ovisacs  absent  ?     Prostates  in  XVII-XX.     Sper- 
mathecae  2  pairs  ;  diverticulum  simple. 

TYPE-LOCALITY.    Lake  St  Clair,  i46°io'E.  42°o5'S.    Map,  13/5. 
MATERIAL  EXAMINED. 

The  single  type-specimen  (NMV  :  Gaga)  is  in  a  very  refractory  condition  and  yields  no  useful 
information. 

REMARKS.  There  is  a  discrepancy  in  Spencer's  account :  genital  markings  are 
paired  in  19/20  according  to  the  text  but  in  18/19  m  the  illustration.  If  the  markings 
were  in  19/20  there  is  a  possibility  that  P.  (P.)  capensis  (from  Table  Cape)  is  a  junior 
synonym.  Both  species  are  perichaetine.  Differences  from  capensis  indicated  by 
Spencer's  account  are  location  of  the  spermathecal  pores  in  db  (not  c),  absence  of 
accessory  genital  markings  in  17/18  and  intestinal  origin  in  XVII  (not  £XVI).  These 
differences,  if  real,  appear  to  warrant  specific  distinction  of  capensis  but  confirmation 
of  the  distinctness  of  the  two  entities  must  await  collection  of  new  material  of  moroea. 

P.  (F.)  evansi  from  the  same  locality  as  moroea  has,  like  the  latter  and  capensis, 
2  pairs  of  spermathecae.  It  differs  from  moroea  in  having  lumbricine  setae,  in  the 
genital  fields  and  in  other  respects  and  conspecificity  is  not  indicated. 


Perionychella  (subgenus  ?)  richea  (Spencer,  1895) 
Fig.  i. 

Perichaeta  richea  Spencer,  1895  :  49-53,  PI.  V,  fig.  49-51. 
Diporochaeta  richea  ;   Jamieson,  igyic  :  83. 

1  =  75  mm,  w  =  3  mm,  s  ?. 

Epilobous  \.  First  dorsal  pore  in  3/4.  Setae  12  on  each  side  anterior  to  clitellum, 
24  half-way  along  the  body.  Clitellum  annular,  XIV- XVI I.  Male  pores  on  papillae 
in  ab.  No  accessory  genital  markings  developed.  Female  pores  on  XIV.  Sper- 
mathecal pores  5  pairs,  on  small  elliptical  patches  at  the  posterior  margins  of  IV- IX, 
in  a. 

Last  hearts  in  XII ;  supra-oesophageal  in  IX-XII.  Gizzards  in  III  and  IV  (?). 
No  true  calciferous  glands  but  oesophagus  in  XI  and  XII  white  and  swollen.  In- 
testinal origin  XVII.  Holonephric.  Holandric ;  seminal  vesicles  racemose,  in 
IX  and  XII.  Metagynous  ;  ovisacs  in  XIV.  Prostates  wide,  tubular,  in  XVII- 
XIX.  Spermathecae  5  pairs  ;  the  diverticulum  very  small. 

TYPE-LOCALITY.  Mount  Olympus,  under  logs  in  the  Beech  forest,  i46°io'E. 
42°05'S.  Map,  13/5. 

MATERIAL  EXAMINED. 

None  available. 

REMARKS.  Reported  location  of  gizzards  in  III  and  IV,  if  accurate,  would  be 
highly  distinctive  but  this  position  is  very  questionable  as  it  is  unknown  elsewhere  in 
the  suborder  Lumbricina.  This  species  must  be  regarded  as  a  species  dubium  as, 


TASMANIAN  EARTHWORMS  259 

apart  from  the  dubious  location  of  the  gizzards,  there  is  nothing  distinctive  in  the 
description  which  would  permit  identification.  The  types  are  no  longer  traceable 
in  the  Spencer  collection  (Jensz  and  Smith  1969). 


Perionychella  (subgenus  ?)  scolecoidea  (Spencer,  1895) 

Fig.  i. 

Perichaeta  scolecoidea  Spencer,  1895  :  51-52,  PI.  V,  fig.  47-51. 
Diporochaeta  scolecoidea  ;  Michaelsen,  1900  :  207  ;   Jamieson,  19710  :  83. 

1  =  25-27  mm,  w  =  5  mm,  s  =  72-77  (lectotype  and  2  paralectotypes). 

Form  very  wide  relative  to  the  short  length.  Prostomium  epilobous  <  £.  First 
dorsal  pore  in  3/4  (Spencer).  Setae  35  on  each  side  in  IV  and  caudally  ;  40-44  on 
each  side  in  XX  ;  no  dorsal  gap  throughout ;  some  ventral  interruption  posteriorly 
but  irregular  and  slight  (paralectotype  i).  Nephropores  and  clitellum  not  visible. 
Male  pores  on  minute  papillae  in  setal  lines  4  in  XVIII.  Accessory  genital  markings 
an  indistinct  transverse  pad  extending  laterally  as  far  as  setal  line  6  in  each  of  inter- 
segments 13/14  and  14/15  (lectotype  and  paralectotype  i ;  none  recorded  in  the  text 
by  Spencer  but  a  median  pad  illustrated  in  19/20).  Female  pores  just  in  front  of  setae 
b  of  XIV.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9,  in  or  slightly  lateral  of  setal 
lines  3  (lectotype)  ;  or  in  s.l.  4  (Spencer). 

Dorsal  vessel  single  ;  last  hearts  in  XII  (confirmation).  Supra-oesophageal  vessel 
in  VIII-XII  (Spencer).  Gizzard  very  large,  but  not  very  thick  walled,  in  V  (con- 
firmation) ;  true  calciferous  glands  absent ;  intestinal  origin  XVIII  (Spencer). 

Holonephric  ;  nephridia  more  coiled  in  II-V  than  elsewhere.  Holandric  ;  seminal 
vesicles  racemose,  in  XII.  Metagynous  (Spencer).  Prostates  racemose,  depressed 
and  lobed,  in  XVIII  (confirmation  from  lectotype).  Penial  setae  absent  (lectotype). 
Spermathecae  2  pairs,  in  VIII  and  IX,  each  with  2  very  small  diverticula  at  the  base 
of  a  fair-sized  sac  (Spencer). 

TYPE-LOCALITY.     King  River  Valley,  approximately   i45°25'E.  42°io'S.     Map, 

I2/I. 

MATERIAL  EXAMINED. 

Lectotype  NMV  :  0290  and  2  paralectotypes,  01422. 

REMARKS.  The  material  examined  was  refractory  and  yielded  little  information. 
The  condition  of  the  nephridial  ducts  was  indeterminable. 


Genus  GRALIOPHILUS  Jamieson,  19716 

DIAGNOSIS.  Combined  male  and  prostatic  pores  a  pair  on  XVIII.  Prostates 
tubular.  Setae  8  per  segment ;  cd  =  o^-i-^bc.  Nephridia  exonephric,  stomate, 
avesiculate  holonephridia  forming  a  single  series  on  each  side  discharging  in  c  or  d 


260  B.    G.   M.    JAMIESON 

or  above  these  ;  sometimes  replaced  in  a  varying  number  of  anterior  segments  by 
coiled  or  tufted  nephridia  with  anteriorly  directed  (always  buccal  ?)  ducts.  Calci- 
ferous  glands  and  intestinal  typhlosole  present  or  absent.  Spermathecae  2  or  less 
commonly  3  pairs  with  extramural  diverticula,  sometimes  with  sperm  chambers  in 
the  walls  but  never  multiloculate. 

TYPE-SPECIES.     Graliophilus  georgei  Jamieson,  19716. 

DISTRIBUTION.  Western  Subregion  of  Australia,  South  Western  faunal  province. 
Eastern  Subregion,  New  South  Wales  (Mt  Kosciusko)  ?,  Tasmania  ?. 

REMARKS.  The  above  generic  diagnosis  is  based  on  Jamieson,  19716.  Gralio- 
philus was  erected  for  Western  Australian  species  which  previously  would  have  been 
ascribable  to  Plutellus  but  clearly  were  not  congeneric  with  the  type-species  of  that 
genus  nor  with  species  of  any  other  genera  erected  during  progressive  revision  of 
Plutellus.  The  two  species  from  New  South  Wales  (G.  montiskosciuskoi  and  G. 
woodi  Jamieson,  I973a),  were  referred  tentatively  to  Graliophilus  but  the  widening  of 
Perionychella  to  include  avesiculate  and  vesiculate,  lumbricine  and  perichaetine 
species,  permits  G.  montiskosciuskoi  to  be  placed  unequivocally  in  Perionychella  as  a 
lumbricine  species  of  the  nominate  subgenus.  G.  woodi  and  the  Tasmanian  species 
G.  tripapillatus  sp.  n.  differ  from  Perionychella  as  currently  denned  in  possessing 
true  typhlosoles.  Typhlosoles  are  present  in  some  Western  Australian  species  of 
Graliophilus  and  woodi  and  tripapillatus  have  therefore  been  referred  to  this  genus, 
with  which  they  conform  in  general  morphology,  though  the  identification  is  made 
with  considerable  hesitation.  It  must  be  noted  that  in  another  Tasmanian  genus, 
Cryptodrilus,  typhlosoles  may  be  present  or  absent  in  closely  related  species  and  that 
separation  of  woodi  and  tripapillatus  from  Perionychella  may,  therefore,  be  unnatural 
as  no  other  significant  features  'are  known  to  exclude  them  from  the  latter  genus. 
Furthermore,  P.  (P.)  capensis  has  a  slight  dorsal  intestinal  ridge  which  might  be 
considered  a  rudimentary  typhlosole. 

This  difficulty  in  generic  identification  of  species  reflects  that  fact  that  distinction 
of  Graliophilus  from  Perionychella  is  rendered  uncertain  by  the  emendment  of 
Perionychella.  While  some  Western  Australian  species  groups,  notably  the  strelitzi- 
group  of  Graliophilus,  are  unquestionably  distinct  from  Perionychella,  the  georgei- 
group,  containing  the  type-species  of  Graliophilus,  conforms  closely  in  morphology 
with  the  emended  Perionychella.  However,  in  Western  Australian  Graliophilus 
species  the  number  of  spermathecae  is  less  (2  or  3  pairs)  than  the  5  pairs  usual  in 
Perionychella  (though  some  species  of  the  latter  have  1-4  pairs)  and  the  avesiculate 
and  totally  lumbricine  condition  is  constant  whereas  the  combination  of  these  charac- 
ters is  known  in  no  Perionychella  species  from  Victoria  or  New  South  Wales  and  in 
only  one,  P.  (P.)  weldborough.' ,  from  Tasmania.  Wallace  (1972)  has  shown  that  species 
since  placed  by  the  author  in  Perionychella  show  only  a  low  phenetic  resemblance  to 
Graliophilus  species  when  affinity  is  assessed  from  large  character-sets  using  computer 
techniques.  It  is  hoped  that  analysis  of  the  data  will  allow  diagnosis  of  each  genus 
from  the  other  without  recourse  to  numerical  methods. 

In  short,  the  following  Tasmanian  species  referred  to  Graliophilus  may  be  a 
typhlosolate  species  of  Perionychella  (Perionychella) . 


TASMANIAN  EARTHWORMS 


261 


sp.p 


sp.p 


sp.p 


XII 


XIII 


XIV— 


1mm 


1mm 


FIG.  17.     Genital  fields.     Graliophilus  tripapillatus ,  holotype  :   A,  spermathecal  field  ; 

B,  male  field. 

Graliophilus  (?)  tripapillatus  sp.  n. 

Fig.  4,  17,  32  A  ;  Plate  i.     Table  3 
1  =  60,  59  mm,  w  =  3-2,  3-0  mm,  s  =  130,  135. 

Prostomium  epilobous  £,  closed.  Dorsal  pores  not  visible.  Setae  8  per  segment, 
in  regular  longitudinal  rows  throughout.  Nephropores  in  c  lines.  Clitellum  annular, 
^XIII-^XVIII.  Male  pores  on  XVIII  in  a  on  dome-shaped  papillae  reaching  from 
b  to  an  approximately  equal  distance  median  to  the  pores.  Accessory  genital  mark- 
ings midventral  unpaired  transverse  pads  in  15/16,  16/17  and  19/20,  extending  later- 
ally into  ab  or  beyond  b  and  longitudinally  to  or  over  the  setal  rows  of  adjacent 
segments  ;  the  pads  increasing  in  size  and  prominence  posteriorly  ;  faintly  indicated 
glandular  and  slightly  tumid  interruptions  of  the  intersegments,  in  17/18  and  18/19. 
Female  pores  paired  on  XIV,  anterior  and  only  slightly  median  to  setae  a.  Sper- 
mathecal pores  3  pairs,  in  6/7-8/9,  on  small  papillae  in  a. 


262 


B.   G.  M.   JAMIESON 


TABLE  3 

Intersetal  distances  in  segment  XII  in  Gmliophilus  tripapillatus 


mm 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

0-9 

0-4 

I-O 

0-6 

3'3 

0-6 

I-O 

0-4 

0-7 

0-4 

0-9 

0-7 

2-7 

0-6 

0-8 

0-4 

i-i 

0-4 

I-O 

0-6 

3-6 

0-6 

I-O 

0-4 

i'3 

0-4 

I-O 

0-6 

3-8 

0-6 

I-O 

0-4 

i-o 

o-5 

0-9 

0-6 

2-8 

0-6 

0-9 

0-4 

standardized  (u  =  100) 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

"'35 

4-80 

11-79 

7-42 

49-61 

7-42 

11-79 

4-80 

10-40 

4'95 

12-87 

9-41 

38-61 

7.92 

10-89 

4-95 

12-55 

4-86 

ii-34 

6-88 

41-30 

6-88 

n-34 

4-86 

13-96 

4'53 

11-32 

6-79 

40-75 

6-79 

11-32 

4'53 

12-44 

5'99 

11-98 

8-29 

36-87 

7-37 

11-98 

5-07 

12-14 

5-03 

11-86 

7.76 

39-63 

7-28 

11-46 

4-84 

2-41 

I-OO 

2-36 

1-54 

7-88 

i-45 

2-28 

0-96 

holotype 
paratype  i 
paratype  2 
paratype  3 
paratype  4 


holotype 
paratype  i 
paratype  2 
paratype  3 
paratype  4 
mean 
interval/afe 


Last  hearts  in  XII  (latero-oesophageal)  ;  supra-oesophageal  moderately  developed 
in  VIII-XIII.  Gizzard  large  and  firm  in  V.  Calciferous  glands  absent ;  oesopha- 
gus internally  rugose  and  with  circumferential  vascular  striae  in  VIII-XVI.  In- 
testinal origin  XVIII ;  deep  dorsal  typhlosole  commencing  in  XX.  Nephridia 
simple  stomate,  avesculate,  exonephric  holonephridia  (funnels  demonstrated  in  fore- 
and  mid-body)  ;  ducts  entering  the  parietes  presetally  in  c  lines,  in  the  forebody 
forming  wide-tubes  but  not  sufficiently  dilated  to  be  termed  bladders ;  slender  in 
the  midbody. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous ;  seminal  vesicles 
racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  in  XIV.  Prostates  much  coiled, 
depressed  tubular ;  vas  deferens  joining  the  sinuous  ectally  widening  duct  shortly 
ectal  of  the  gland.  Penial  setae  very  slender,  tip  bifid,  chelate,  the  two  branches 
terminally  contiguous,  a  coarse  tooth,  almost  large  enough  to  be  considered  a  bifurca- 
tion, on  one  branch  ;  ornamentation  absent ;  length  ?  mm,  general  width  of  shaft 
=  3  /mi  (holotype).  Spermathecae  3  pairs  ;  diverticulum  (inseminated)  single, 
clavate,  uniloculate. 

Genital  markings.  Of  the  15  type-specimens  (holotype,  paratypes  1-14)  only  the 
holotype  and  paratype  i  have  the  genital  marking  in  15/16.  Otherwise  the  genital 
fields  are  invariable  in  all  specimens,  with  median  genital  marking  in  each  of  inter- 
segments 16/17  and  19/20  and,  less  strongly  indicated,  in  17/18  and  18/19. 

MATERIAL  EXAMINED. 

Tarraleah,  over  pipeline,  I46°25'E.  42°2o'S.  Map,  13/4,  Dr  J.  L.  Hickman,  27  May  1954  - 
holotype  TM  :  K3I3  ;  paratypes  2-6  TM  :  1^314-318  ;  paratypes  i,  7-9  BM(NH)  :  1973.2.1-4  ; 
paratypes  10-12  AM  :  W5203-52O5  ;  paratypes  13  and  14  BJ  :  T23-24- 

REMARKS.  The  genital  field,  with  its  arrangement  of  three  genital  markings, 
diagnoses  this  species. 


TASMANIAN  EARTHWORMS  263 

Genus  PINGUIDRILUS  gen.  nov. 

DIAGNOSIS.  Setae  8  per  segment.  Combined  male  and  prostatic  pores  a  pair  on 
XVIII.  Prostates  a  single  pair,  tubular.  Gizzard  in  V  and  VI ;  extramural  calci- 
ferous  glands  and  intestinal  typhlosole  absent.  Nephridia  vesiculate  holonephridia 
discharging  irregularly  in  c  to  near  middorsum.  Spermathecae  adiverticulate. 

DESCRIPTION.  Large  ;  terrestrial.  Dorsal  pores  present.  Setae  8  per  segment. 
Nephropores  erratically  in  c  to  near  the  middorsal  line.  Clitellum  long  (>  9 
segments),  including  the  male  pores.  Male  pores  (combined  pores  of  the  vasa 
deferentia  and  the  single  pair  of  prostates)  on  XVIII.  Female  pores  paired  on  XIV. 
Spermathecal  pores  5  pairs,  the  last  in  8/9.  Accessory  genital  markings  present  in 
the  vicinity  of  the  male  pores. 

Last  hearts  in  XIII,  latero-oesophageal  hearts,  in  X-XIII,  with  connectives  to 
dorsal  and  the  supra-oesophageal  vessel.  Gizzard  large,  in  V-VI.  Extramural 
calciferous  glands  absent ;  intestinal  origin  XIX  ;  typhlosole  absent.  Nephridia 
simple,  exonephric  holonephridia  with  large  ectal  bladders.  Holandric  ;  gymnor- 
chous  ;  seminal  vesicles  in  IX  and  XII.  Metagynous  ;  ovisacs  absent.  Prostates 
slightly  lobulated  tubes.  Penial  setae  absent.  Spermathecae  lacking  diverticula. 

DISTRIBUTION.     Tasmania.     (Monotypic.) 
TYPE-SPECIES.    Notoscolex  tasmanianus  Fletcher,  i888b. 

REMARKS.  Pinguidrilus  retains  the  primitive  holonephric  condition  (two  neph- 
ridia  per  segment)  seen  elsewhere  in  Tasmania  only  in  Rhododrilus  (Acanthodrilinae) 
and  in  Perionychella  and  a  doubtful  member  of  each  of  the  genera  Graliophilus  and 
Woodwardiella  (Megascolecinae).  The  italicized  features  in  the  above  diagnosis 
distinguish  it  from  all  these  genera.  Its  closest  phyletic  relationship  is  presumably 
with  the  similarly  vesiculate  Perionychella  (V  esiculodrilus]  but  the  absence  of  sper- 
mathecal  diverticula  is  a  very  rare  condition  in  the  Megascolecidae  (with  the  excep- 
tion of  the  Ocnerodrilinae,  in  which  diverticula  are  only  exceptionally  present)  and, 
with  the  unusual  arrangement  of  the  nephropores,  suggests  a  rather  isolated  position 
for  the  genus.  This  irregular  disposition  of  the  nephropores  is  also  seen  in  the 
Nearctic  plutelloid  genus  Argilophilus,  which  shows  strong  morphological  affinities 
with  Pinguidrilus,  though  avesiculate  and  possessing  spermathecal  diverticula 
albeit  little  more  than  sperm  chambers  in  the  wall  of  the  spermathecal  duct. 

Pinguidrilus  tasmanianus  (Fletcher,  i888b) 
Fig.  4,  256,  326 

Notoscolex  tasmanianus  Fletcher,  i888b  :  607-611. 

Megascolides  tasmanianus  ;   Spencer,  1895  :  33- 

Plutellus  tasmanianus  ;   Michaelsen,  1900  :  175  ;   Jamieson,  19710  :  88. 

1  =  ?,  w  =  23  mm,  s  =  ?. 

Circular  in  cross  section,  segments  I  and  II  simple  ;  thereafter,  in  the  forebody, 
strongly  triannulate.  Prostomium  epilobous,  closed,  \  ;  it  and  the  peristomium 


264  B.   G.  M.   JAMIESON 

with  numerous  longitudinal  grooves.  First  dorsal  pore  in  10/11,  minute.  Setae 
8  per  segment  but  c  and  d  only  sporadically  visible  ;  setae  a  and  b  absent  in  XVIII. 
Nephropores  (see  nephridia,  below).  Clitellum  annular,  strongly  protuberant, 
embracing  XIV-^XXIII ;  dorsal  pores  absent,  intersegmental  furrows  retained 
only  ventrally  ;  ventral  setal  couples  visible  ;  nephropores  not  visible.  Male  pores 
distinct  but  minute  apertures  in  ab  on  small  white  papillae  which  occupy  the  setal 
annulus  of  XVIII.  Genital  markings  7  median  transverse  pads,  in  intersegmental 
furrows  14/15-20/21,  each  bisected  by  the  furrow,  longitudinally  occupying  the  pos- 
terior and  anterior  annuli  of  the  adjacent  segments,  and  extending  laterally  beyond 
setal  lines  b  ;  those  in  17/18  and  18/19  not  as  extensive  transversely  as  the  others  and 
only  shortly  overreaching  b  lines.  Female  pores  a  pair  of  minute  orifices  on  small, 
low  papillae,  on  XIV,  very  shortly  anteromedian  of  setae  a,  on  the  setal  annulus. 
Spermathecal  pores  on  5  pairs  of  small,  round  papillae,  in  4/5-8/9,  in  a  lines. 

Septa  progressively  thickened  from  3/4  to  12/13  ;  3/4  and  4/5  obscured  by  tendons 
running  to  the  next  posterior  septa  ;  5/6  strong  ;  12/13  immensely  thick  ;  13/14 
moderately  thick  ;  the  remainder  thin.  Dorsal  blood  vessel  single  ;  continuous  onto 
the  pharynx.  Dorsoventral  commissural  vessels  in  V-XIII ;  those  in  X-XIII 
forming  4  pairs  of  latero-oesophageal  hearts  ;  those  in  IX  anteriorly  progressively 
more  slender  and  dorsoventral  only.  Each  latero-oesophageal  heart  receiving  a 
slender  connective  from  the  dorsal  vessel  and  a  thicker  connective  from  the  median 
supra-oesophageal  vessel  immediately  behind  the  junction  with  the  latter  of  a  pair  of 
vessels  from  the  oesophageal  wall.  Supra-oesophageal  beginning  (very  thin) 
anteriorly  in  VIII  and  ending  posteriorly  as  the  connectives  to  the  hearts  in  XIII,  the 
vessel  well  defined  and  separate  from  the  oesophagus  throughout  and  posteriorly 
almost  as  thick  as  the  dorsal  vessel ;  receiving  vessels  from  the  oesophagus  throughout 
its  course.  Gizzard  large  and  muscular  but  fairly  easily  compressible,  in  V  and  VI, 
septum  5/6  being  inserted  on  its  equator  where  the  commissures  of  V  join  the  dorsal 
vessel.  Oesophagus  in  VII  long;  coiled  and  muscular  ;  in  VIII-XI  vascular,  though 
not  conspicuously,  and  not  dilated  ;  in  XII  and  XIII  segmentally  dilated  and  with 
numerous  circumferential  vascular  striae  ;  in  XIV- XVI 1 1  very  wide,  extremely 
thin  walled  and  vascular,  giving  paired  vessels  directly  to  the  dorsal  vessel. 
Intestine  commencing  in  XIX  ;  no  typhylosole  detectable. 

Nephridia  :  nephropores  erratically  positioned  (described  by  Fletcher  as  an 
irregularly  sinuous  series  varying  from  c  lines  to  near  the  middorsal  line)  ;  their 
position  sometimes  corresponds  with  setal  loci  but  these  also  are  erratic.  In  the 
intestinal  region  nephridia  are  stomate,  vesiculate  holonephridia  without  supple- 
mentary loops  ;  vesicles  are  large,  in  two  wide  sections  with  a  narrow  section  be- 
tween ;  a  slight  angular  projection  which  could  be  regarded  as  a  diverticulum  extends 
laterally  beyond  the  position  of  opening  ;  no  change  in  structure  occurs  at  the  hind 
end  ;  anteriorly  nephridial  structure  is  similar  but  the  vesicle  is  not  distinctly  divided 
into  two  parts  (Wallace,  personal  communication).  Testes  ?  sperm  masses  and  large 
funnels  free  in  X  and  XI.  Seminal  vesicles  large  and  racemose  in  IX  and  XII ; 
approximately  equisized.  Ovaries  paddle-shaped,  with  many  rows  of  minute 
oocytes,  and  small  funnels  in  XIII.  Prostates  restricted  to  XVIII,  slender  tor- 
tuous tubes  with  slightly  lobulated  surfaces ;  the  slender  ducts  receiving  the  vasa 


TASMANIAN   EARTHWORMS  265 

deferentia  near  the  junction  with  the  glandular  portions.  Penial  setae  absent. 
Spermathecae  5  pairs  of  simple  pouches  tapering  to  the  pores  but  lacking  distinctly 
differentiated  ducts  ;  diver ticula  absent. 

TYPE-LOCALITY.    Thomas'  Plains,  N.E.  Tasmania. 
MATERIAL  EXAMINED. 

A  single  fragmenting  clitellate  specimen  labelled  ' Megascolides  tasmanicus  [sic]  Fletcher  jar 
2',  Baldwin  Spencer  collection,  ex  Melbourne  University  Zoology  Department,  May  1968. 

REMARKS.  The  above  description  agrees  closely  with  that  of  Fletcher  and,  as  the 
material  of  Fletcher  species  in  the  Baldwin  Spencer  collection  must  be  considered 
type  material,  the  specimen  examined  is  here  designated  the  lectotype  NMV  :  G2H2. 
The  following  additional  data  are  given  by  Fletcher  :  lengths  (contracted)  200-250 
mm,  width  19-24  mm  ;  about  200  segments.  Setal  lines  d  sinuous  ;  aa  :  ab  :  be  :  cd 
=  4:2:4:  3-8.  The  first  dorsal  pore  was  observed  in  12/13,  and  the  clitellum  on 
^XIII-XXII ;  the  gizzard  in  V  ;  the  oesophagus  dilated  and  vascular  in  VIII-XVI 
and  especially  in  XIII-XVI,  showing  effervesence  in  acid  in  all  but  the  first  segment. 


Genus  WOODWARDIELLA  Stephenson,  1925 

DIAGNOSIS.  Combined  male  and  prostatic  pores  a  pair  on  XVIII.  Prostates 
racemose.  Setae  8  per  segment ;  cd  =  or  slightly  >  be.  Nephridia  exonephric, 
stomate,  avesiculate  holonephridia  forming  a  single  series  on  each  side  in  c  lines  but 
(always  ?)  replaced  in  anterior  segments  by  tufts  discharging  into  the  buccal  cavity 
or  exonephrically  in  d  lines.  Calcif  erous  glands  absent ;  typhlosole  weakly  developed 
or  absent.  Spermathecae  2  pairs,  diverticulum  simple,  not  multiloculate.  (Based 
on  Jamieson  (1970,  19716).) 

TYPE-SPECIES.     Woodwardia  callichaeta  Michaelsen,  i9O7a. 
DISTRIBUTION.     South-western  Australia.     Tasmania  ?     Victoria  ? 

REMARKS.  Differences  between  Woodwardiella  and  the  older  genus  Perionychella 
require  augmentation.  The  prostates  in  Perionychella  are  tubular  or,  if  externally 
racemose,  retain  an  axial  lumen  whereas  those  of  Woodwardiella  have  a  typical 
Pheretima-type  prostate  with  branching  internal  ducts  (Michaelsen,  1916).  Al- 
though this  difference  is  allowed  in  Cryptodrilus,  a  genus  in  which  species  with  the 
two  alternative  conditions  of  the  prostates  are  assignable  to  the  same  genus  by  virtue 
of  the  peculiar  excretory  system  which  they  share,  it  warrants  separation  of  Wood- 
wardiella and  Perionychella  pending  further  investigation  of  the  former  little  known 
genus.  Tubular  prostates  are  all  that  is  certainly  known  to  distinguish  Graliophilus 
from  the  older  Woodwardiella  and  further  investigation  of  the  justification  of  retain- 
ing Graliophilus  as  a  separate  genus  from  Woodwardiella  or  Perionychella  is  also 
required. 

The  following  Tasmanian  species  was  transferred  to  Woodwardiella  by  Jamieson 
(i97ic)  from  the  artificial  congeries  Plutellus,  in  which  Michaelsen  (1900)  had  placed 
it  on  the  then  justified  grounds  that  it  had  8  setae  per  segment,  holonephridia  and 
apparently  racemose  prostates.  As  no  material  of  it  is  available  for  the  present 

16 


266  B.   G.   M.   JAMIESON 

study  it  is  retained  in  Woodwardiella  though  it  appears  likely  that  its  prostates  will  be 
found  to  be  tubulo-racemose  and  that  it  will  be  assignable  to  Perionychella. 

Woodwardiella  ?  tessellatus  (Spencer,  1895) 
Fig.  4 

Cryptodrilus  tesselatus  (sic)  Spencer,  1896  :  40-41,  PI.  II,  fig.  16-18. 
Plutellus  tessellatus  ;   Michaelsen,  1900  :  170-171. 
Woodwardiella  ?  tesselatus  ;   Jamieson,  I97ic  :  92. 

1  =  25  mm,  s  =  65. 

Prostomium  slightly  epilobous.  First  dorsal  pore  in  5/6.  Setae  8  per  segment, 
the  dorsal  row  very  irregular  and  setae  d  absent  in  some  segments  ;  c  regular  except 
caudally.  Clitellum  annular,  XIII -XVII,  with  midventral  continuation  in  bb  in 
XVIII  and  XIX.  Male  pores  on  small  papillae  in  XVIII,  in  ab.  Accessory  genital 
markings  2  pairs  of  small  elliptical  patches  in  12/13  and  13/14  in  ab.  Female  pores 
anteromedian  to  setae  a,  at  \aa,  in  XIV.  Spermathecal  pores  2  pairs,  in  7/8  and 
8/9,  in  ab. 

Last  hearts  in  XII.  Gizzard  in  V  ;  true  calciferous  glands  absent ;  intestinal 
origin  XVII.  Holonephric.  Holandric  ;  seminal  vesicles  saccular,  in  XII.  Meta- 
gynous.  Prostates  small  flattened  lobes,  in  XVIII.  Spermathecae  2  pairs,  in  VIII 
and  IX  ;  diverticulum  simple,  less  than  half  the  length  of  the  ampulla. 

TYPE-LOCALITY.  Mount  Olympus,  Lake  St  Clair,  in  damp  soil  under  logs,  and 
amongst  decaying  leaves  in  beech  forest.  Map,  13/5. 

MATERIAL  EXAMINED. 

None  available. 

Genus  CRYPTODRILUS  Fletcher,  i887a  emend. 

DIAGNOSIS.  A  pair  of  combined  male  and  prostatic  pores  on  the  seventeenth 
setigerous  segment,  normally  XVIII.  Setae  8  per  segment.  Meronephric  ;  neph- 
ridia  typically  vesiculate,  frequently  avesiculate,  throughout,  not  very  numerous  on 
each  side,  some  or  all  discharging  in  the  setal  lines  ;  the  medianmost  nephridium 
usually  (always  ?  )  with  preseptal  funnel.  Typhlosole  present  or  absent.  Prostates 
tubular,  tubuloracemose,  or  racemose. 

DESCRIPTION.  Moderate-sized  earthworms  with  less  than  200  segments.  Pro- 
stomium zygolobous  to  tanylobous.  Dorsal  pores  present  or  absent.  Setae  in  8 
regular  longitudinal  lines,  commencing  on  II  (or,  C.  mediocris,  on  the  product  of 
union  of  the  first  and  second  metameres).  Ventral  setal  couples  widely  paired 
(aa  =  i-5~3a6)  ;  setae  of  the  dorsal  couples  (cd)  widely  separated,  always  further 
apart  than  those  of  the  ventral  couple  (cd  =  2-4^6)  and  usually  a  greater  interval, 
sometimes  smaller,  than  the  interval  between  the  couples  of  a  side  (be)  ;  dorsal 
median  intersetal  distance  (dd)  ==  0-2-0-4  of  the  circumference  (u).  Setae  a  and  b 
absent  in  the  segment  bearing  the  male  pores  ;  usually  replaced  by  penial  setae. 
Nephropores  (conspicuous  or  not)  at  least  3  on  each  side  in  some  segments  and 


TASMANIAN   EARTHWORMS  267 

reaching  (C.  polynephricus)  as  many  as  10  per  side  ;  the  pores  associated  with  some  or 
all  four  of  the  setal  lines  where  the  number  per  side  is  4  or  less  ;  increase  in  number 
intersetal  and  setal.  Clitellum  annular,  commencing  in  XI I -XIV  and  including  or 
shortly  preceding  the  segment  bearing  the  male  pores.  A  pair  of  combined  male  and 
prostatic  pores  on  XVIII  (or  XVII  where  intersegment  1/2  is  suppressed),  in  b  or 
median  of  this.  Accessory  genital  markings  present.  Female  pores  a  pair,  or  un- 
paired midventral,  in  XIV,  inconspicuous,  though  sometimes  in  a  common  glandular 
field.  Spermathecal  pores  2  or  3  pairs,  the  last  pair  in  intersegmental  furrow  8/9. 

Some  preclitellar  septa  strongly  thickened.  Gizzard  rudimentary  to  well 
developed,  in  V.  Definite  calciferous  glands  absent  but  oesophagus  in  some  seg- 
ments vascularized,  often  dilated,  and  internally  rugose  or  lamellate.  Intestinal 
origin  XVI,  XVII  or  XVIII ;  typhlosole  present  or  absent.  Dorsal  blood  vessel 
(always  ?)  continued  onto  the  pharynx  ;  dorso ventral  commissurals  commencing  in 
V  or  further  posteriorly  ;  last  hearts  in  XII  or  XIII,  those  in  X-XIII  forming  latero- 
oesophageal  hearts  with  connectives  from  the  supra-oesophageal  vessel  and  in  some 
species  from  the  dorsal  vessel.  Supra-oesophageal  vessel  not  extending  more  than  a 
segment  or  two  in  front  of  the  latero-oesophageal  hearts.  Subneural  vessel  absent. 
Nephridia  3~ca.  15  exonephric  nephridia  on  each  side,  typically  vesiculate  but  fre- 
quently avesiculate,  some  or  all  discharging  in  the  setal  lines  ;  the  medianmost 
nephridium  in  the  intestinal  or  at  least  caudal  region  stomate  in  most  if  not  all  species 
and  frequently  enlarged  as  a  megameronephridium  ;  posterior  nephridia  sometimes 
sending  ducts  posteriorly  through  several  segments  or  sometimes  with  the  stomate 
nephridia  contributing  to  a  longitudinal  excretory  duct  (ureter)  which  appears  to 
discharge  at  junction  of  rectum  and  body  wall ;  rarely  (C.  enteronephricus)  with  duct 
of  each  stomate  megameronephridium  entering  the  intestine.  Avesiculate  exo- 
nephric or  enteronephric  tufts  present  or  absent  in  anterior  segments.  Holandric  ; 
gymnorchous  ;  seminal  vesicles  in  2  or  more  of  segments  IX,  X,  XI  and  XII. 
Metagynous  ;  ovisacs  present  or  absent.  Prostates  one  pair,  racemose,  with  branched 
internal  and  external  ducts,  or  tubuloracemose  (lobulated  but  with  single  central 
lumen  which  (always  ?)  has  side  branches)  ;  vasa  deferentia  joining  the  ectal  end  of 
the  gland  or  the  prostate  duct  shortly  ectal  of  this.  Spermathecae  discharging 
anteriorly  in  their  segments  ;  the  duct  with  one  or  two  uniloculate  diverticula. 

TYPE-SPECIES.     Cryptodrilus  rusticus  Fletcher,  iSSya. 

DISTRIBUTION.  Eastern  Subregion  of  Australia,  Southern  faunal  province : 
New  South  Wales  and  Victoria.  Tasmania. 

REMARKS.  The  tribe  Perionychini  was  established  by  Jamieson  (iQ7ia)  for  those 
Megascolecinae  which  were  holonephric,  or  had  meronephridia  in  a  varying  number 
of  segments  anterior  to  holonephridia,  and  which  lacked  intestinal  enteronephry. 
Demonstration  of  more  than  two  vesiculate  nephridia  per  segment  in  Cryptodrilus 
(vide  Jamieson  iQ72b)  posed  the  problem  that  this  genus  could  not  be  placed  in  any 
one  of  the  three  megascolecine  tribes  (Perionychini,  Dichogastrini  and  Megascolecini) 
recognized  by  the  author  but  studies  of  general  morphology,  and  a  numerical  (taxono- 
metric)  investigation  of  setal  ratios,  suggested  that  the  affinities  of  Cryptodrilus  lay 
with  the  Perionychini.  An  evolutionary  pathway  for  origin  of  the  multiple-bladder 


268  B.   G.   M.    JAMIESON 

condition  from  the  holonephric  vesiculate  condition  exemplified  by  Plutellus  hetero- 
porus,  the  'heteropore'  condition,  was  suggested.  It  was  pointed  out,  and  is  still 
maintained,  that  recognition  of  a  tribe  Cryptodrilini  to  include  Cryptodrilus  deserves 
consideration.  In  the  present  work  Cryptodrilus  is  included  in  the  Perionychini 
because  of  its  close  morphological  and  presumed  evolutionary  links  with  this  tribe, 
the  nephridial  bladders,  which  are  unknown  in  the  Dichogastrini  and  Megascolecini, 
being  an  especially  significant  indication  of  affinity.  Pending  the  possibility  of 
erection  of  a  tribe  Cryptodrilini,  then,  the  definition  of  the  Perionychini  must  be 
enlarged  to  include  species  with  multiple  bladders  in  a  segment  or,  in  other  words,  all 
megascolecine  species  with  nephridial  bladders,  irrespective  of  whether  holonephric 
or  meronephric,  in  addition  to  holonephric  avesiculate  species. 

Some  species  of  Cryptodrilus,  which  have  clear  affinities  (sometimes  at  the  infra- 
specific  level)  with  vesiculate  species,  lack  nephridial  bladders.  They  have  median 
stomate  nephridia  in  posterior  segments,  as  do  some  vesiculate  species,  and  are 
presumed  to  represent  an  acquisition  of  the  dichogastrin  condition  of  nephridia 
independently  of  the  true  Dichogastrini.  This  view  is  preferred  to  the  alternative 
that  they  represent  a  survival  of  populations  transitional  between  the  Perionychini 
and  Dichogastrini  as  the  numerical  study  indicated  that,  at  least  in  setal  ratios,  the 
Dichogastrini  are  distinct  from  the  Perionychini.  This  distinctness  might  have  been 
developed  subsequent  to  origin  of  Dichogastrini  from  populations  of  which  Tas- 
manian  perionychines  are  representative  but  variation  from  the  vesiculate  to  the 
avesiculate  condition  infraspecifically,  or  at  least  in  very  closely  related  subgeneric 
taxa  (vide  C.  polynephricus),  indicates  that  loss  of  vesicles  is  currently  occurring  in 
Tasmanian  perionychines.  The  definition  of  the  Perionychini  must,  therefore,  be 
further  extended  to  allow  inclusion  of  forms  which  have  acquired  the  dichogastrin 
condition  but  have  clear  relationships  with  perionychin  species.  In  addition,  an 
avesiculate  species,  C.  enteronephricus,  with  intestinal  enteronephry  (a  condition 
formerly  restricted  to  the  Megascolecini)  which  is  unquestionably  related  to  vesiculate 
species  of  Cryptodrilus,  must  be  included  within  this  genus  in  the  Perionychini.  Thus 
the  definition  of  the  Perionychini  embraces  species  which  would  be  referable  to  the 
Dichogastrini  or  Megascolecini  on  the  anatomy  of  the  excretory  system  but  which 
show  evident  relationships  with  Perionychini. 

Such  widening  of  the  definition  of  discrete  groups  so  that  they  are  no  longer  totally 
interdistinguishable  has  become  accepted  in  modern  taxonomy  and  is  an  expected 
extension  of  the  concept  of  polythetic  groups,  definable  by  no  one  single  character. 

The  genus  Cryptodrilus,  as  emended  above  to  include  additional  evidence  from 
Tasmanian  species,  has  now  become  a  polythetic  grouping  as  the  single  formerly 
diagnostic  character,  multiple  nephridial  bladders  in  a  segment,  no  longer  applies  to 
all  species.  Inclusion  of  avesiculate  species  appears  justified  from  a  consideration  of 
general  anatomy.  Such  a  variation,  from  vesiculate  to  avesiculate  species  is  in- 
significant relative  to  the  remarkable  variation  further  discussed  in  the  account  of  C. 
polynephricus,  from  the  exonephric  to  the  enteronephric  condition  in  posterior  seg- 
ments. 

Jamieson  (igyaa)  recognized  three  species  groups  in  Cryptodrilus.  The  Tasmanian 
species  C.  avesiculatus,  C.  brunyensis,  C.  enteronephricus  and  C.  polynephricus  (with 


TASMANIAN   EARTHWORMS  269 

C.  albertisi)  show  clear  affinities  with  the  mediocris  group  and  C.  simsoni  and  less 
certainly  C.  dartnalli  probably  also  are  placeable  in  it.  It  is  not  proposed,  however, 
to  redefine  the  group  as  the  emended  definition  would  be  so  wide  as  to  be  poorly 
defined  from  the  rusticus  and  fastigatus  groups  and  would  have  little  operational 
value.  This  is  not  to  deny  that  at  least  fastigatus,  with  its  externally  branched 
prostates,  is  a  very  unusual  species.  If  further  species  with  fastigatus  prostates  were 
found  it  would  not  be  unreasonable  to  recognize  a  separate  genus  for  such  species, 
the  name  Trinephrus  being  available  for  it. 

CHECK  LIST  OF  SPECIES  OF  Cryptodrilus 
Tasmanian  species 

C.  albertisi  (Cognetti,  1910)  New  combination  (junior  synonym  of  C.  polynephri- 

cus  ?) 

C.  avesiculatus  sp.  n. 
C.  brunyensis  sp.  n. 
C.  dartnalli  sp.  n. 
C.  enteronephricus  sp.  n. 
C.  officeri  Spencer,  1895 
C.  polynephricus  Spencer,  1895 
C.  simsoni  Spencer,  1895 

Other  species 

New  South  Wales 

C.  fastigatus  Fletcher,  iSSga 

C.  mediocris  Fletcher,  i88ga 

C.  rusticus  Fletcher, 

C.  tenuis  Fletcher, 

Victoria 

C.  dubius  Spencer,  i892a  (junior  synonym  of  C.  fastigatus) 

KEY  TO  TASMANIAN  SPECIES  OF  CRYPTODRILUS* 

1  3  pairs  of  spermathecae      .......          C.  officeri  Spencer,  1895 

-  2  pairs  of  spermathecae       ...........  2 

2  Gizzard  large,  firm.     Intestine  commencing  in  XVI        ......  3 

-  Gizzard  small,  flaccid.     Intestine  commencing  in  XVII  or  XVIII   ....  6 

3  Caudally  with  median  stomate  nephridium  on  each  side  discharging  into  the  intestine. 

Fig.  20  .........       C.  enteronephricus  sp.  n. 

-  No  caudal  nephridia  discharging  into  the  intestine  unless  anally      ....  4 

4  2  nephridia  present  caudally  in  a  line  on  each  side  :    i  astomate  and  discharging 

exonephrically,  the  other  stomate  and  contributing  to  a  longitudinal  collecting 

duct  (ureter)  ............  5 

-  Only  one  nephridium  present  caudally  in  a  line  on  each  side,  astomate  and  exonephric. 

No  ureter  present.     Fig.  i8B C.  avesiculatus  sp.  n. 

-  Excluding  C.  albertisi  (q.v.) 


ayo 


B.    G.   M.    JAMIESON 


1mm 


sp.p 


sp.p 


g.m 


FIG.  1  8.     Genital  fields.     A,  Hickmaniella  opisthogaster,  holotype.     B,  Cryptodrilus 

avesiculatus,  holotype. 


5  No  postsetal  genital  markings  in  preclitellar  segments.     Fig.  igA   .       C.  brunyensis  sp.  n. 

-  Postsetal  genital  markings  present  in  preclitellar  segments.     Fig.  2iA,  B,  C 

C.  polynephricus  Spencer,  1895 

6  Dorsal  pores  present.    Intestine  commencing  in  XVIII.    Fig.  25  A    C.  sitnsoni  Spencer,  1895 

-  Dorsal  pores  absent.     Intestine  commencing  in  XVII.     Fig.  196    .  C.  dartnalli  sp.  n. 


Cryptodrilus  albertisi  (Cognetti,  1910) 
Fig.  2 

Megascolides  albertisi  Cognetti,  1910  :  329-331,  Fig.  3-6. 
1  =  68  mm,  w  =  4  mm,  s  =  107. 

Prostomium   tanylobous.     Dorsal   pores   not   observed.     Ventral   setae   closely 
paired,  dorsal  setae  distant,  aa  <  be  <  ^throughout.     InV-XVIII,  aa  =  ab  +  be  ; 


TASMANIAN   EARTHWORMS  271 


ab  >  be  ;  cd  =  2bc  ;  dd  =  I'^cd  =  o-^u.  Behind  XVIII  setal  lines,  with  the  excep- 
tion of  a,  displaced  dorsally  so  that,  in  XXX,  aa  slightly  >  ab,  but  appreciably 
<  ab  +  be  ;  ab  slightly  <  be  ;  cd  >  2bc  ;  dd  =  aa  =  0-14^.  Clitellum  not  deve- 
loped. Male  pores  not  externally  visible.  Accessory  genital  markings  3  pairs  of 
whitish,  slightly  tumid  papillae  posteriorly  in  segments  IX,  X  and  XI  in  b  lines. 
Female  pores  a  pair  anteriorly  in  XIV,  midway  between  setal  lines  a  and  the  ventral 
midline.  Spermathecal  pores  2  pairs  of  short  transverse  slits  in  7/8  and  8/9,  shortly 
lateral  of  a  lines. 

Last  hearts  in  XIII.  Gizzard  extending  from  mid  V  to  mid  VI  ;  calciferous 
glands  absent  ;  intestinal  origin  XVI.  Nephridia  'diffuse'  ;  from  approximately 
segment  XXX,  those  of  the  most  ventral  pair,  in  b  lines,  are  much  larger  than  the 
others.  Holandric  ;  gymnorchous  ;  seminal  vesicles  small,  in  IX,  larger  in  XII, 
non-racemose.  Prostates  one  pair,  in  XVIII-XX,  the  glandular  part  oblong  with 
corrugated  surface  ;  the  S-shaped  muscular  duct  penetrating  the  parietes  in, 
apparently,  b.  Penial  setae  800  £im  long,  30  /u,m  wide,  slightly  curved,  ending  in  a 
point  which  is  not  very  fine  ;  the  ectal  half  ornamented  with  small  scale-like  projec- 
tions with  minutely  dentate  margins.  Metagynous.  Spermathecae  2  pairs,  in 
VIII  and  IX  ;  ampulla  transversely  compressed  with  slightly  lobulated  margin  with 
stout  duct  of  equal  length  into  which,  a  short  distance  from  the  external  pore,  opens 
a  digitiform  diverticulum. 

TYPE-LOCALITY.    Mt  Wellington.    Map,  14/4. 

MATERIAL  EXAMINED. 

None  available. 

REMARKS.  Cognetti's  description  is  questionably  adequate  for  identification  of 
any  material  with  this  species.  Although  nephridial  bladders  and  distribution  of 
nephridia  are  not  described,  the  distribution  of  accessory  genital  markings  strongly 
suggests  that  C.  albertisi  is  identical  with,  and  therefore,  a  junior  synonym  of, 
C.  polynephricus  Spencer,  1895. 

Cryptodrilus  avesiculatus  sp.  n. 
Fig.  2,  i8B,  3iA  ;  Plates  2  and  3.     Table  4 
1  =  81,  69  mm,  w  =  3-8,  4-0  mm,  s  =  116,  118. 

Prostomium  epilobous  f  ,  open.  Dorsal  pores  absent.  Setae  8  per  segment,  in 
regular  longitudinal  rows  throughout.  Nephropores  not  visible.  Clitellum  annular, 
JXIII-XVII.  Male  and  separate  penisetal  pores  on  dome-shaped  papillae  in  b  of 
XVIII.  Accessory  genital  markings  2  or  3  pairs  of  eyelike  markings  in  17/18 
(holotype),  18/19  and  19/20  (holotype,  paratype  i)  in  a  lines  ;  postsetal  paired  oval 
markings  extending  over  the  posterior  borders  of  their  segments  in  VII  and  VIII 
containing  the  spermathecal  pores  (holotype)  >and  in  IX  (holotype,  paratype  i)  ; 
also  a  faintly  indicated  paired  postsetal  patch  in  and  ventral  to  a  in  VII  and  VIII 
(holotype)  .  Female  pores  shortly  anteromedian  to  setae  a  of  XIV.  Spermathecal 
pores  2  pairs,  in  7/8  and  8/9,  on  minute  papillae  in  ab. 


272 


B.   G.  M.   JAMIESON 


TABLE  4 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  avesiculatus 


holotype 
paratype  3 
paratype  4 
paratype  5 


holotype 
paratype  3 
paratype  4 
paratype  5 
mean 
interval/aft 


aa  ab  be  cd  dd  dc  cb  ba 

1-6  0-7  1-4  1-5  3-5  1-5  1-4  0-7 

i-i  0-6  1-2  1-4  2-4  1-4  i-i  0-6 

i-i  0-6  i-i  1-3  2-3  1-3  i-o  0-6 

i '4  0-6  i-i  1-3  2-1  1-3  i-o  0-6 

standardized  (u  =  100) 


aa 

06 

be 

cd 

dd 

dc 

cfc 

ba 

13-07 

5-68 

11-36 

12-50 

28-69 

11-93 

11-36 

5-40 

n-33 

6-37 

12-04 

14-51 

24-43 

13-81 

"•33 

6-02 

11-97 

6'73 

11-22 

14-21 

24-68 

I3-46 

10-85 

6'73 

14-96 

6-36 

"•59 

13-84 

22-44 

13-84 

10-47 

6-36 

12-83 

6-29 

"•55 

I3-77 

25-06 

13-26 

11-00 

6-13 

2-04 

i-oo 

1-84 

2-19 

3-98 

2-II 

i-75 

0-97 

Last  hearts  in  XII  (latero-oesophageal).  Supra-oesophageal  present.  Gizzard 
large,  in  V.  Extramural  calciferous  glands  absent  but  oesophagus  vascularized  and 
moniliform  in  IX-XIII  (holotype,  paratype  i)  and  slightly  in  VIII  (paratype  i). 
Intestinal  origin  XVI ;  typhlosole  absent.  Nephridia  (holotype)  :  large  (exone- 
phric  ?)  tufts  in  III  and  IV  ;  succeeding  segments  with  a  few  large  avesiculate  integu- 
mentary micromeronephridia  on  each  side.  Caudally  4  exonephric,  avesiculate 
nephridia  on  each  side  discharging  in  front  of  setae  a,  b,  c  and  d ;  the  dorsal  three 
astomate,  the  medianmost  nephridium,  discharging  in  a  with  a  large  preseptal  funnel 
and  greatly  enlarged  to  form  a  megameronephridium  ;  one  nephridium  only  at  the  a 
locus,  the  megameronephridium ;  no  ureter.  Holandric  (funnels  iridescent)  ; 
gymnorchous  ;  seminal  vesicles  compact,  racemose  in  IX  and  XII.  Metagynous  ; 
ovisacs  absent.  Prostates  flattened,  lobulated  but  linear  winding  through  XVII- 
XX  ;  with  narrow  central  lumen.  Vasa  deferentia  joining  the  ectal  end  of  the  gland. 
Penial  setae  moderately  stout,  strongly  curved,  widened  basally,  the  smooth  delicate 
pointed  tip  upturned  ;  the  ectal  region  of  the  shaft,  with  the  exception  of  the  vicinity 
of  the  tip,  ornamented  by  a  series  of  coarsely  serrated,  scarcely  protuberant  trans- 
verse but  appreciably  oblique  jagged  markings  each  of  which  occupies  approximately 
one-quarter  to  three-quarters  of  the  width  of  the  seta  ;  length  of  functional  seta  = 
1-3  mm,  general  width  of  shaft  =  23  /urn  (holotype).  Spermathecae  2  pairs ; 
diverticulum  single,  subspherical,  in  the  axis  of  the  duct ;  ampulla  a  lateral  appen- 
dage. 

Genital  markings.  (Holotype,  paratypes  1-7.)  Genital  markings  in  IX  are 
restricted  to  the  Port  Davey  specimens.  In  the  new  Harbour  specimens  a  distinct 
oval  genital  marking  is  developed  in  front  of  each  spermathecal  pore,  on  VII  and 
VIII  ;  there  are  only  suggestions  of  these  in  the  Port  Davey  specimens.  All 
specimens,  from  both  localities,  have  paired  eye-like  marking  is  18/19  and  19/20 
but  only  the  holotype  and  paratype  2  have  these  markings  in  17/18  and  in  only 
paratype  3  are  they  present  (median  to  a]  in  20/21. 


TASMANIAN  EARTHWORMS 


273 


sp.p 


sp.p 


VIII 


XVIII 

XIX- 

XX- 

XXI- 


linfn 


FIG.  19.     Genital  fields.     A,  Cryptodrilus  brunyensis,  holotype.     B,  C.  dartnalli,  holotype. 

C,  C.  simsoni,  8/1,  specimen  2. 

MATERIAL  EXAMINED. 

Kelly's  Basin,  Port  Davey,  I45°55'E.  43°2o'S.  Map,  17/1,  Tasmanian  Biological  Survey  : 
Jiy,  Mr  C.  D.  King,  Jan  1940  -  holotype  TM  :  K3ig  ;  paratype  i  BM(NH)  :  1973.2.5  ;  paratype 
2  TM  :  K320.  New  Harbour,  i46°7o'E.  43°3o'S.  Map,  18/5,  Mr  C.  D.  King  -  paratypes  3 
and  4  TM  :  K32I-322  ;  paratypes  5  and  6  AM  :  W5206-5207  ;  paratype  7  BJ  :  T25. 

REMARKS.  C.  avesiculatus  is  closely  similar  to  C.  enteronephricus,  C.  polynephricus 
or  C.  brunyensis.  It  differs  from  C.  enteronephricus  in  lacking  caudal  enteronephry, 
and  from  C.  polynephricus  and  C.  brunyensis  in  lacking  a  ureter. 


Cryptodrilus  brunyensis  sp.  n. 
Fig.  2,  igA,  316,  C  ;  Plates  4-7.     Table  5 


1  =  48  mm,  71  mm,  w  =  3-0  mm,  s  =  83,  124. 

Prostomium  epilobous  \,  open.  Dorsal  pores  absent.  Setae  8  per  segment  in 
regular  longitudinal  rows  throughout.  Nephropores  not  visible.  Clitellum  annular 
XIII-XVII.  Male  pores  in  ab,  nearer  a,  in  XVIII  on  subcircular  papillae  which 
extend  posteriorly  over  18/19  slightly  into  XIX  and  anteriorly  reach  the  hind  margins 
of  a  pair  of  oval  genital  markings  in  ab  at  17/18.  Two  penial  setae,  projecting  from 
separate  follicles,  median  to  each  pore.  Two  additional  pairs  of  genital  markings 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

ii-45 

6-17 

12-33 

15-86 

18-94 

15-86 

13-66 

5-73 

13-03 

6-90 

12-64 

I5-7I 

16-48 

I5-7I 

12-64 

6-90 

12-24 

6-54 

12-49 

15-79 

17-71 

15-79 

13-15 

6-32 

1-87 

I  -00 

1-91 

2-41 

2-71 

2-41 

2-OI 

0-97 

274  B-   G-   M-    JAMIESON 

present  in  19/20  and  20/21  median  to  a,  the  anterior  pair  fused  medianly  (holotype)  ; 
only  the  conjoined  pair  in  19/20  present  in  paratype  i.  Female  pore  unpaired, 
midventral,  between  the  setal  arc  and  anterior  margin  of  XIV.  Spermathecal 
pores  2  pairs,  in  7/8  and  8/9,  in  a  lines,  each  surrounded  by  a  glandular  field  ;  the 
fields  in  7/8  (holotype)  or  in  7/8  and  8/9  (paratype)  medianly  united  and  suggestions 
of  presetal  markings  present  in  b  of  IX  in  paratype  i. 

TABLE  5 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  brunyensis 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

holotype  0-9  0-5  i-o  1-3  1-5  1-3  i-i  0-5 

paratype  I  1-2  0-6  1-2  1-4  1-5  1-4  1-2  0-6 

standardized  (u  =  100) 

aa  ab  be 

holotype 
paratype  i 
mean 
interval/a& 

Dorsal  blood  vessel  continuous  onto  the  pharynx.  Last  hearts  in  XII ;  hearts  in 
X-XII  latero-oesophageal,  each  with  a  connective  to  the  supra-oesophageal  vessel 
but  none  to  the  dorsal  vessel,  supra-oesophageal  ending  at  |XIII,  well  developed. 
Gizzard  firm  and  very  large,  in  -V.  Calciferous  glands  absent  but  oesophagus  intern- 
ally lamellate  in  VIII-XIII.  Intestinal  origin  XVI ;  dorsal  typhlosole  a  low  but 
distinctly  developed  lamina  from  XVIII  but  continued  as  a  rudiment  to  XVI 
(holotype,  paratype).  Nephridia  :  loose  aggregations  of  nephridia  in  II  and  III 
send  several  ducts  anteriorly  to  discharge  at  the  anterior  margin  of  the  segment ; 
pharyngeal  tufts  in  IV- VI,  accompanied  by  exonephric  meronephridia  ;  in  VII  and 
VIII  with  a  meronephridium  discharging  above  d,  at  d,  cd,  c,  b  and  several,  of  which 
some  form  a  small  tuft,  at  a  ;  this  condition  persisting  in  succeeding  segments 
(holotype)  ;  in  the  hindbody  with  a  micromeronephridium  discharging  in  the  4  setal 
lines  a,  b,  c  and  d  and  in  addition  a  stomate  megameronephridium  contributing  its 
duct  to  a  poorly  developed  longitudinal  collecting  duct  (ureter)  (holotype,  paratype) 
which  ends  posteriorly  at  the  junction  of  rectum  and  body  wall  and  therefore  appears 
to  discharge  into  the  anal  aperture  on  each  side  ;  all  nephridia  avesiculate  and  only 
the  megameronephridia  with  funnels  (holotype).  In  paratype  the  ureter  is  well 
developed  but  is  not  certainly  detectable  at  the  caudal  extremity  where  in  a  there  is 
only  a  single  (astomate  ?)  nephridium  ;  those  in  b,  c  and  d  persisting. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles 
racemose  in  XI  and  XII.  Metagynous  ;  ovisacs  present.  Prostates  broad,  flat- 
tened, linear,  lobulated  (tubuloracemose) ,  in  XVIII-XXI  (holotype,  paratype), 
dissected  to  demonstrate  main  central  internal  duct  with  numerous  more  slender 
lateral  branches  (holotype).  Vasa  deferentia  on  each  side  winding  before  joining  the 


TASMANIAN  EARTHWORMS  275 

prostate  duct  at  its  junction  with  the  gland.  Penial  seta  moderately  stout,  strongly 
curved,  tapering  to  a  narrow  blunt  or  pointed  tip  ;  the  ectal  region  of  the  shaft,  with 
the  exception  of  the  vicinity  of  the  tip,  ornamented  by  numerous  groups  of  small, 
irregular,  conjoined  teeth,  the  groups  alternating  along  the  seta,  each  containing  only 
a  few  teeth  but  considerably  notching  the  setal  surface  ;  length  of  a  functional  seta  = 
1-9  mm  ;  general  width  of  shaft  =  20  pm  (paratype  i).  Spermathecae  (not  insemi- 
nated) 2  pairs,  each  with  long  convoluted  duct  and  long  clavate-digitiform  diver- 
ticulum. 

MATERIAL  EXAMINED. 

Rainforest  at  summit  of  Mt  Mangana,  Bruny  Is.,  147°I5'E.  43°25'S.  Map,  19/1,  Mr  A.  J. 
Dartnall,  10  Apr  1971  -  holotype  TM  :  1X323  ;  paratype  BM(NH)  :  1973.2.6. 

REMARKS.  C.  brunyensis  is  morphologically  close  to  C.  enteronephricus,  C. 
avesiculatus  and  C.  polynephricus.  It  differs  from  C.  enteronephricus  in  lacking 
caudal  enteronephry,  from  C.  avesiculatus  in  possessing  a  ureter  and  from  C.  poly- 
nephricus in  lacking  preclitellar  genital  markings. 


Cryptodrilus  dartnalli  sp.  n. 

Fig.  2,  196,  3iD  ;   Plates  8-12.     Table  6 

1  =  85  +  ,  109  mm,  w  =  3-7,  3-5  mm,  s  =  107  + ,  143  (holotype,  posterior  amputee  ; 
paratype  2). 

Prostomium  epilobous  \,  closed,  continued  posteriorly  as  parallel  grooves,  almost 
giving  a  tanylobous  condition.  Dorsal  pores  absent.  Setae  8  per  segment,  in 
regular  longitudinal  rows  throughout.  Nephropores  sporadically  visible  presetally. 
Clitellum  not  developed  (holotype)  or  strongly  protuberant,  though  weak  in  aa,  in 
XIV-XVII  with  some  dorsal  clitellar  modification  to  |XVIII  and  perhaps  |XIII 
(paratype  2).  Male  pores  small  slits  on  XVIII  in  a  on  ovoid  papillae  which  extend 
laterally  as  far  as  b  and  well  median  of  a.  Glandular  tumescence  extending  from  setal 
arc  of  XVII  to  shortly  behind  18/19,  reaching  mid  be  at  17/18,  and  including  the  male 
porophores  (holotype),  similar  in  paratype  I  in  which  a  pair  of  faint  eye-like  genital 
markings  is  present  on  the  tumescence  in  17/18  in  b  ;  indistinctly  developed  in 
paratype  2  and  other  New  Harbour  specimens  and  obscured  in  these  by  insinking  of 
the  male  field.  Female  pores  a  pair  very  shortly  anteromedian  of  setae  a  on  XIV. 
Spermathecal  pores  2  pairs  of  small  slits  on  small  but  distinct  papillae,  in  7/8  and 
8/9,  in  a. 

TABLE  6 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  dartnalli 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

i-i  0-6  1-3  i-2  4-5  1-2  1-3  0-6 

i-i  0-7  1-5  1-4  4-8  1-5  1-6  0-7 

1-2  0-8  1-6  1-4  5-8  1-3  1-6  0-7 


276 
Table  6  (cont.) 

B.   G.   M.   JAMIESON 

standardized  (u  =  100) 

aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

holotype 

9-15 

4-88 

10-98 

10-37 

38-41 

10-37 

10-98 

4-88 

paratype  i 

8-59 

5-37 

11-28 

IO-2O 

35-98 

11-28 

11-81 

5'37 

paratype  2 

8-16 

5-28 

11-04 

9-60 

39-60 

9-12 

11-04 

4-80 

mean 

8-63 

5-18 

II-IO 

10-06 

38-00 

10-26 

11-28 

5-02 

interval/a& 

1-67 

I  -00 

2-14 

1-94 

7-34 

1-98 

2-18 

0-97 

sp.p 


1mm 


FIG.  20.     Genital  fields.     Cryptodrilus  enteronephricus  :   A,  holotype  ;   B,  paratype  18  ; 

C,  paratype  3. 


TASMANIAN   EARTHWORMS  277 

Last  hearts  in  XIII  (latero-oesophageal ;  connectives  to  dorsal  vessel  not  demon- 
strable). Supra-oesophageal  in  VII-XIII,  well  developed.  Gizzard  vestigial,  in  V, 
but  musculature  thicker  than  elsewhere.  Extramural  calciferous  glands  absent ; 
oesophagus  with  circumferential  vascular  striae  and  internal  papilliform  rugae,  in 
VII-XIII  (holotype),  XIV  (paratype)  ;  chloragogenous  and  narrower  in  XIV 
(holotype),  XV  (paratype  2) -XVI,  though  with  internal  rugae  in  XIV  in  holotype  ; 
intestinal  origin  XVII  ;  typhlosole  absent.  Nephridia  :  in  the  forebody,  astomate, 
avesiculate,  exonephric  meronephridia  commencing  in  II,  few  in  number,  all  (?) 
associated  with  setal  lines  a,  b,  c  and  d  ;  tufts  absent.  Caudally  in  the  four  setal 
lines,  or  in  a-c  and  above  d,  the  median-most  nephridium  with  a  preseptal  funnel. 
Holandric  (funnels  iridescent  in  X  and  XI) ;  gymnorchous  ;  seminal  vesicles  spheroi- 
dal, in  IX  and  XII.  Metagynous ;  ovisacs  absent  (holotype),  but  small  sacs 
(ovisacs  ?)  in  XIV  in  paratype  2.  Prostates  flattened  tubular,  tortuous,  in  XVIII 
and  XIX  ;  two  vasa  deferentia  joining  the  junction  of  each  gland  with  its  sinuous 
muscular  duct  (holotype)  or  joining  the  ectal  region  of  the  gland  (paratype  2).  Penial 
setae  moderately  stout,  gently  curved,  tapering  to  a  narrow  point ;  the  ectal  region 
of  the  shaft,  with  the  exception  of  the  vicinity  of  the  tip,  ornamented  by  oblique 
laterally  sometimes  interrupted  circlets  of  irregular  coarse  teeth  which  are  directed 
anteriorly  but  may  be  considerably  elevated  at  their  apices  from  the  setal  surface  ; 
the  circlets  approximately  7  /zm  apart  (holotype,  paratype  8)  ;  length  of  functional 
seta  o-8-i-o  mm  ;  general  width  of  shaft  22, 17  ju,m  (holotype,  paratype  8).  Sperma- 
thecae  2  pairs.  Diverticulum  single,  digitiform  uniloculate  ;  shorter  relative  to 
the  ampulla  in  paratype  2  compared  with  the  holotype. 

MATERIAL  EXAMINED. 

Melaleuca  Inlet,  Port  Davey,  I45°55'E.  43°2o'S.  Map,  17/1,  Mr  A.  J.  Dartnall,  Dec  1967  - 
holotype  TM  :  Kaao,  paratype  i  TM  :  1^324.  New  Harbour,  i46°io'E.  43°3o'S.  Map,  18/5, 
Mr  C.  D.  King,  27  Jan  1938  -  paratypes  2  and  3  BM(NH)  :  1973.2.7-8  ;  paratypes  4-6 
AM  :  W52o8-52io  ;  paratypes  7-8  BJ  :  T26-27. 

REMARKS.  The  genital  field  of  C.  dartnalli  is  diagnostic  but  the  species  shows 
close  affinities  with  C.  simsoni.  It  differs  from  the  latter,  among  other  respects, 
in  lacking  dorsal  pores  and  in  origin  of  the  intestine  one  segment  further  forward, 
in  XVII. 

Cryptodrilus  enteronephricus  sp.  n. 
Fig.  2,  20,  3iE-G  ;  Plates  13-19.     Table  7 
1  =  124,  no  mm,  w  =  4-6,  4-0  mm,  s  =  162,  168. 

Prostomium  epilobous  f  with  narrow  open  dorsal  tongue.  Dorsal  pores  absent. 
Setae  8  per  segment  in  regular  longitudinal  rows  throughout.  Nephropores  not,  or 
only  sporadically,  visible.  Clitellum  annular,  |XIII-XVIII  (paratype  i)  or  XVII, 
with  weak  extension  to  |XVIII  (holotype).  Male  pores  small  transverse  slits,  not 
on  porophores,  in  ab  of  XVIII,  nearer  a  than  b  (holotype)  or  (paratype  i)  nearer  b 
than  a,  with  two  penisetal  pores  median  to  each  pore,  the  three  pores  of  a  side  on  a 
small  indefinite  papilla.  Paired  accessory  genital  markings  at  the  posterior  borders 


278  B.   G.   M.    JAMIESON 

of  IX  and  X  in  ab,  intersegmental  in  16/17  m  a  and  I7/I8  and  18/19  m  a^  (holotype, 
paratype  i),  and  intersegmental  in  19/20  in  and  median  to  ab  (holotype)  ;  median 
markings  occupying  aa  in  14/15  and  15/16  (holotype,  paratype  i).  Female  pores 
anteromedian  to  setae  a  of  XIV,  at  approximately  \aa.  Spermathecal  pores  2  pairs, 
in  7/8  and  8/9,  on  small  distinctly  protuberant  papillae  in  ab,  shortly  lateral  of  a. 

TABLE  7 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  enter onephricus 


holotype 
paratype  i 
paratype  3 
paratype  4 
paratype  6 
paratype  14 
paratype  15 


aa 

ab      j 

be     i 

:d 

dd 

dc      < 

cb 

ba 

•o 

o-5 

•3 

•3 

2-0 

i-3 

•3 

o-5 

•3 

0-6 

•5 

•8 

2-7 

1-7 

•5 

0-6 

•2 

o-5 

•3 

•3 

1-8 

1-3 

•3 

o-5 

•2 

0-6 

•9 

•6 

2-6 

1-6 

•o 

0-6 

•3 

0-6 

•7 

•8 

2-5 

1-8 

•7 

0-6 

•0 

0-6 

•3 

•6 

2-0 

i'5 

:4 

0-6 

•7 

0-8 

•5 

•9 

2-6 

1-9 

•5 

I-O 

standardized  (u  =  100) 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

10-89 

5-06 

14-40 

14-01 

21-79 

14-01 

14-79 

5-06 

11-03 

5-07 

13-11 

I5-50 

22-65 

14-30 

13-11 

5-07 

12-55 

4'94 

I4-45 

14-45 

19-77 

M'45 

14-45 

4-94 

10-09 

4-90 

I5-56 

13-26 

21-61 

13-26 

16-43 

4-90 

10-88 

5-00 

13-82 

15-29 

20-88 

15-29 

13-82 

5-00 

10-15 

5-60 

12-60 

I5-75 

19-95 

I5-40 

13-65 

5-60 

12-69 

6-48 

11-61 

14-85 

19-98 

I4-58 

n-34 

7-29 

11-18 

5-29 

I3-65 

I4-73 

20-95 

14-47 

13-94 

5-4i 

2-II 

I'OO 

2-58 

2-78 

3-96 

2-74 

2-64 

1-02 

holotype 
paratype  i 
paratype  3 
paratype  4 
paratype  6 
paratype  14 
paratype  15 
mean 
interval/a& 

Last  hearts  in  XIII  (latero-oesophageal  with  supraoesophageal  but  no  dorsal 
connectives  in  X-XIII).  Commissurals  in  IX  anteriorly,  slender  and  dorsoventral, 
lacking  supra-oesophageal  connectives.  Supra-oesophageal  vessel  in  VIII-XIV, 
well  developed. 

Gizzard  large  and  firm,  with  anterior  rim,  in  V.  Calciferous  glands  absent. 
Oesophagus  vascularized  in  VIII-XIV.  Intestinal  origin  XVI ;  a  deep  laminar 
dorsal  typhlosole  present,  rapidly  increasing  in  depth  from  XVIII,  continued  as  a 
very  low  ridge  or  thickening  into  XVI  ;  muscular  thickening  and  caeca  absent. 
Nephridia  :  large  tufts  in  II-V  send  sheaves  of  forwardly  directed  ducts  to  the 
buccal  cavity  and  pharynx,  those  in  II  discharging  at  the  junction  of  buccal  cavity 
and  body  wall,  those  in  III  entering  the  buccal  cavity  and  those  in  IV  and  V  joining 
the  pharynx  behind-  the  brain.  Succeeding  segments  with  several  astomate, 
avesiculate,  exonephric  parietal  micromeronephridia  on  each  side.  Caudally  with 
4  exonephric,  astomate,  micromeronephridia  on  each  side  discharging  in  a,  b,  c  and 
d  lines  and  in  addition  a  very  large  stomate  megameronephridium  which  has  a  large 
preseptal  funnel  near  the  ventral  nerve  cord  and  loops  far  laterally  (to  approximately 
c  line)  before  turning  medianly  to  send  a  duct  to  the  ventrolateral  surface  of  the 


TASMANIAN  EARTHWORMS 


279 


intestine,  i.e.  enteronephric.  All  5  nephridia  on  each  side  persisting  to  the  caudal 
extremity.  No  ureter  present. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles 
racemose,  in  IX,  X  and  XII.  Metagynous  ;  small  ovisacs  in  XIV.  Prostates  linear, 
depressed,  winding  in  XVIII-XXI ;  much  lobulated  but  with  a  narrow  axial  lumen 
(tubuloracemose)  but  that  on  the  right  side  in  the  holotype  a  broad  flattened  lobe 
in  XVI 1 1- XX  and  not  resolvable  into  a  linear  form  ;  vasa  deferentia  joining  gland 
near  its  junction  with  the  sinuous  muscular  duct.  Penial  setae  moderately  stout, 
gently  curved,  tapering  ectally  to  a  simple,  moderately  sharp  (paratype  i)  or  (arte- 
fact ?)  truncated  tip  (paratype  14)  ;  the  ectal  region  of  the  shaft,  with  the  exception 
of  the  vicinity  of  the  tip  ornamented  by  about  20  irregular  serrated  circlets  approxi- 
mately 6  )u,m  apart ;  the  serrations  forming  jagged  lines  with,  here  and  there, 
individual  narrow  pointed  teeth  recognizable.  The  circlets  complete  or  interrupted 
along  the  lateral  aspect  of  the  seta,  if  interrupted  the  two  halves  tending  to  alternate 
along  the  seta.  Length  1-05,  0-81  mm  ;  general  width  of  shaft  =  28  /*,  22  (para- 
types  i,  14).  Spermathecae  2  pairs,  with  ovoid  ampulla  and  approximately  equally 
long  duct  which  is  joined  at  midlength  by  an  ovoid  unstalked  diverticulum. 

Genital  markings.  The  following  conclusions  are  drawn  from  examination 
of  the  holotype  and  eight  fully  clitellate  paratypes  :  paired  genital  markings  are 
always  present  posteriorly  in  X  but  may  be  absent  (paratypes  4, 5)  in  IX ;  the  greatest 
extent  of  genital  markings  occurs  in  Pio  in  which,  in  addition  to  the  markings  in  IX 
and  X,  there  are  median  markings  in  13/14  and  14/15  and  paired  markings  in  each  of 
intersegmental  furrows  15/16-20/21 ;  markings  are  present  in  all  specimens  in  each 
of  intersegments  15/16-18/19.  Those  in  15/16  are  usually  median  as  are  those  in 
14/15  if  present ;  those  in  13/14  (seen  in  only  two  specimens)  are  median  or  paired  ; 
all  other  markings  are  paired  though  there  may  be  some  tendency  to  unite  medianly. 
Paired  markings  in  IX  and  X  are  virtually  in  line  with  the  spermathecal  pores,  in  ab 
usually  nearer  a. 


Distribution  of  accessory  genital  markings 
in  Cryptodrihis  enteronephricus 


H 


Pi 


P2 


P5 


P6 


Pio 


Total 


IX 

P 

P              P 

P 

P 

P 

P                 7 

X 

P 

P              P 

P             P 

P 

P 

P 

P                 9 

13/14 

P 

m                2 

14/15 

m 

m 

P 

m 

m 

m                6 

15/16 

m 

m            m 

P             Pm 

pm 

m 

m 

P                 9 

16/17 

pm 

P             P 

P             Pm 

P 

P 

pm 

P                  9 

17/18 

P 

P             P 

P             P 

P 

P 

P 

P                  9 

18/19 

P 

P             P 

P             P 

P 

P 

P 

P                  9 

19/20 

P 

P 

P             P 

P 

P                  6 

2O/2I 

P 

P             P 

P 

P                 5 

m  =  median  ; 

p  =  paired  ;   pm  =  paired  tending  to  fuse  medianly. 

H  =  holotype  ; 

P  =  paratype. 

Other   variations.     Seven   specimens,    including   the   holotype,    were   examined 
internally.     Constant  features  included  :  location  of  gizzard  in  V  ;  intestinal  origin 


28o  B.   G.   M.   JAMIESON 

in  XVI ;  large  typhlosole  ;  enteronephric  megameronephridia  caudally  and  absence 
of  bladders  ;  seminal  vesicles  in  IX,  X  and  XII ;  presence  of  penial  setae  and  2  pairs 
of  spermathecae.  A  major  variation  was  location  of  last  hearts  in  XII  in  paratypes 
3,  4  and  5,  in  contrast  with  location  in  XIII  in  the  holotype  and  paratypes  i,  6  and 
10.  Such  variation  is  highly  exceptional  intraspecifically  but  there  appear  to  be  no 
grounds  for  regarding  the  individuals  with  hearts  in  XII  as  specifically  distinct. 
Ovisacs  were  seen  only  in  paratypes  i  and  5. 

MATERIAL  EXAMINED. 

Dee  Bridge,  Lyell  Highway,  I46°35'E.  42°i5'S.  Map,  13/7,  Dr  J.  L.  Hickman,  24  May  1954  ~ 
holotype  TM  :  1^325.  Tarraleah,  Lyell  Highway,  I46°25'E.  42°2o'S.  Map,  13/4,  Dr  J.  L. 
Hickman,  22  May  1954  -  paratypes  4  and  5  AM  :  ^5211-5212  ;  paratypes  18  and  19  BJ  :  T34- 
35.  Tarraleah,  over  pipeline,  Dr  J.  L.  Hickman,  27  May  1954  ~  paratypes  11-15  BJ  :  T28-32. 
Marlborough  Highway  (near  Bronte),  i46°3o'E.  42°io'S.  Map,  13/6,  Dr  J.  L.  Hickman, 
26  May  1954  -  paratype  J  BM(NH)  :  1973.2.9  ;  paratype  2  BJ  :  T33  ;  paratypes  6-10  TM  : 
K326-33O.  Lyell  Highway,  5  miles  from  Bronte  towards  Hobart,  I46°35'E.  42°i5'S.  Map, 
13/7,  Dr  J.  L.  Hickman,  24  May  1954  -paratype  3  BM(NH)  :  1973.2.10. 

REMARKS.  C.  enteronephricus  appears  to  be  related  to  C.  polynephricus,  and 
especially  to  the  subspecies  urethrae,  but  differs  notably  from  that  species  (q.v.)  in 
having  enteronephric  intestinal  nephridia.  The  development  of  enteronephry 
represents  a  major  departure  from  perionychin  organization  and  the  attainment  of  a 
condition  previously  known  only  in  the  distinct  tribe  Megascolecini  which,  in  sens, 
strict.,  is  otherwise  represented  in  Tasmania  only  by  Oreoscolex. 


Cryptodrilus  ?  officeri  Spencer,  1895 
Fig.  2 

Cryptodrilus  officeri  Spencer,  1895  :  44-45,  PI.  Ill,  fig.  28-30. 
Trinephrus  officeri  ;   Michaelsen,  1900  :  186-187. 
Notoscolex  officeri ;   Jamieson,  i97ic  :  79. 
Cryptodrilus  officeri  ;   Jamieson,  19725  :  166. 

1  =  44  mm,  w  =  6  mm,  s  ? 

Prostomium  epilobous  f .  First  dorsal  pore  4/5.  Setae  8  per  segment,  irregular 
caudally.  Clitellum  annular,  XIV-XVII.  Male  pores  on  papillae  on  XVIII,  in  b. 
Accessory  genital  markings  paired  elliptical  patches  in  15/16  and  16/17  in  b,  and  in 
19/20  in  ab.  Female  pores  presetal  and  close  together  in  XIV.  Spermathecal  pores 
3  pairs,  in  6/7-8/9,  shortly  above  b. 

Last  hearts  in  XII,  no  continuous  supra-oesophageal.  Gizzard  well  marked, 
segment  ?  (no  distinct  septa  in  front  of  7/8)  ;  true  calciferous  glands  absent  but 
vascular  swellings  in  XIII-XV  ;  intestinal  origin  XVII.  Nephridia  ;  3  tufts  on 
each  side.  Holandric ;  seminal  vesicles  racemose  in  IX  and  XII.  Metagynous. 
Spermathecae  3  pairs,  in  VII,  VIII  and  IX  ;  diverticulum  a  group  of  finger-like 
processes,  the  ampulla  long  and  irregular  in  outline. 

TYPE-LOCALITY.     King  River  Valley.     Map,  12/1. 


TASMANIAN   EARTHWORMS 


281 


17 


282  B.  G.  M.  JAMIESON 

MATERIAL  EXAMINED. 

None  available. 

REMARKS.  Spencer's  statement  that  the  nephridia  are  like  those  of  C.  fastigatus 
and  C.  dubius  necessitates  placing  officer i  in  Cryptodrilus  but  inclusion  in  this  genus  is 
questionable.  The  multiloculate  spermathecal  diverticulum  raises  the  possibility 
that  it  is  an  Oreoscolex,  the  3  pairs  of  spermathecae  and  the  configuration  of  the 
genital  field  being  reminiscent  of  0.  sexthecatus.  The  latter  species  is  distinguished, 
so  far  as  the  inadequate  description  of  C.  officer  i  indicates,  by  the  more  ventral 
position  of  the  spermathecal  pores  (in  b  lines).  New  material  of  C.  officeri  is  required 
if  its  status  is  to  be  settled. 


Cryptodrilus  polynephricus  Spencer,  1895 

Cryptodrilus  polynephricus  Spencer,  1895  :  35~36  PI-  I>  ng-  4.  5.  6. 
1  =  98-330  mm,  w  (midclitellar)  =  4-5-12  mm,  s  =  127-175. 

Prostomium  epilobous  |,  f  to  tanylobous,  sometimes  canaliculate.  Dorsal  pores 
absent.  Setae  8  per  segment,  in  regular  longitudinal  rows  throughout ;  a  and  b 
absent  on  XVIII.  Nephropores  minute,  only  sporadically  if  at  all  visible.  Clitellum 
annular,  XIII-XVII.  Male  pores  on  XVIII  in  ab,  not  usually  on  papillae  ;  2 
separate  penisetal  follicles  usually  visible  median  to  each  male  pore.  Accessory 
genital  markings  ;  an  anterior  series  of  3  pairs  of  elliptical  markings  posteriorly  in  IX, 
X  and  XI,  or  at  the  posterior  intersegment  of  each  of  these,  in  ab,  or  median  of  a,  or 
some  or  all  of  these  markings  absent,  or  additional  markings  present  at  the  posterior 
margins  of  VII,  VIII  and  XII.  A  posterior  series  of  paired  or  median  transverse 
markings  present  in  or  immediately  anterior  to  some  of  intersegments  12/13-21/22, 
more  commonly  in  17/18-19/20.  Female  pores  in  XIV,  paired  anteromedian  of  a,  or 
unpaired  midventral  and  presetal.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9,  in  or 
slightly  lateral  of  a. 

Last  hearts  in  XIII  (exceptionally,  in  some  specimens  of  urethrae,  in  XII)  those  in 
X-XIII  latero-oesophageal,  with  connectives  from  the  supra-oesophageal  vessel 
only.  Gizzard  large,  strongly  muscular,  with  anterior  rim,  in  V.  Extramural 
calciferous  glands  absent.  Intestinal  origin  XVI ;  typhlosole  absent,  rudimentary 
or  deep  laminar  ;  muscular  thickening  and  caeca  absent.  Nephridia  :  several  pairs 
of  meronephric  tufts  discharging  into  the  foregut ;  exonephric  micromeronephridia 
present  in  most  segments  on  each  side  in  the  four  setal  rows,  usually  with  additional 
intervening  nephridia,  caudally  with  the  medianmost  nephridium  enlarged  as  a 
megameronephridium,  with  preseptal  funnel,  the  duct  of  which  contributes  to  a 
longitudinal  excretory  duct  (ureter)  which  apparently  discharges  into  the  ureter ; 
nephridial  bladders  present  or  absent,  if  present  number  in  a  segment  ranging  from 
4  to  10  per  side.  Testes  and  iridescent  sperm  funnels  in  X  and  XI  ;  gymnorchous  ; 
seminal  vesicles  in  IX  and  XII,  or  XI  and  XII  or  rarely  IX,  XI  and  XII.  Prostates 
tubuloracemose,  extending  through  2  or  more  segments.  Penial  setae  long  and  stout, 
when  functional  0-7-4  mm  long,  with  the  general  width  of  the  shaft  23-45  //,m. 


TASMANIAN   EARTHWORMS  283 

Metagynous  ;    ovisacs  present  or  absent.     Spermathecae  2  pairs,  in  VIII  and  IX, 
diverticulum  single,  digitiform,  uniloculate  with  or  without  a  basal  swelling. 

TYPE-LOCALITY.     Mt  Wellington.     Map,  14/4.     Parattah.     Map,  14/1. 
DISTRIBUTION.     See  subspecies. 

REMARKS.  Two  subspecies  are  here  recognized  and  described  below.  Where  a 
subspecific  character  does  not  vary  from  the  specific  description  it  is  not  repeated  in 
the  subspecific  account. 


sp.p 


VI 


sp.p 


1mm 


FIG.  22.     Genital  fields.     Cryptodrilus  polynephrictts  polynephricus  :   A,  specimen  7 

B,  specimen  24. 


284  B.   G.   M.   JAMIESON 

Cryptodrilus  polynephricus  polynephricus  Spencer,  1895 
Fig.  2,  2iA,  22,  24A,  3iH  ;  Plates  20-31.     Table  8 

Cryptodrilus  polynephricus  Spencer,  1895  :  35-36,  PL  i,  fig.  4,  5,  6. 

Trinephrus  polynephricus  ;   Michaelsen,  1900  :  185-186. 

Cryptodrilus  polynephricus  ;   Jamieson,  19720  :  169-172,  Fig.  6D,  7E,  F,  G. 

1  =  98-200  mm  (44  specimens),  the  extreme  specimens  with  midclitellar  widths  of 
4-5  and  8  mm,  and  segmental  counts  of  127  and  158,  respectively. 

Male  pores  not  usually  on  papillae,  sometimes  on  oval  papillae  which  extend  beyond 
a  and  b  and  occupy  the  posterior  three-fourths  of  the  segment.  Accessory  genital 
markings  :  characteristically  with  3  pairs  of  anterior  genital  markings  (raised  or 
depressed  glandular  ellipses)  posteriorly  in  IX,  X  and  XI,  in  ab,  in  IX  in  or  shortly 
median  to  b  ;  in  X  slightly  median  to  b  or  at  approximately  mid  be  ;  in  XI  at  mid 
ab  or  nearer  but  not  in  a,  i.e.  lateral  of  the  line  of  the  spermathecal  pores  and  conver- 
gent posteriorly. 

TABLE  8 

Intersetal  distances  in  segment  XII  in  14  specimens*  of 
Cryptodrilus  polynephricus  polynephricus 

mm 
aa  ab  be  cd  dd  dc  cb  ba 

maximum  2-3  0-9  2-0  4-1  4-4  4-0  1-9  0-9 

minimum  i-i  0-6  1-3  1-5  2-1  1-4  i-i  0-3 

mean  1-8  0-7  1-6  2-8  3-2  2-7  1-6  0-7 

standard  deviation  0-3  o-i  0-2  0-9  0-7  0-8  0-2  o-i 

standardized  (u  =  100) 
aa  ab  be  cd  dd  dc  cb  ba 

maximum                           I3'45          6'9°        J4'94        20-86  24-41  20-50  13*64  0-36 

minimum                           10-63          3'54          9-24        13-03  18-16  13-22  8-43  3-54 

mean                                   12-19          5-10        11-15        17-89  20-79  17-27  10-69  4'93 

standard  deviation             1-20          0-99          1-63          2-43  1-69  2-35  1-73  0-88 

interval/afc                           2-39          i-oo          2-19          3-51  4-08  3-39  2-10  0-97 
*  Specimens  i,  4,  5,  7,  10,  12,  14,  18,  24,  26,  32,  34,  37  and  38. 

Some  individuals  (14/4,  27  Oct  1955  ;  13  Nov  1954)  lacking  markings  in  IX  and 
with  those  in  X  and  XI  in  or  median  of  a.  Posterior  genital  markings  in  or  immedi- 
ately anterior  to  intersegmental  furrows  :  median  at  14/15,  occasionally  ;  paired  or 
median  at  15/16,  commonly  ;  paired  or  median  at  16/17,  17/18  and  18/19,  almost  in- 
variably ;  paired  but  usually  medianly  conjoined  in  19/20,  usually,  in  20/21, 
commonly,  and  in  21/22  rarely  ;  the  paired  markings  centred  from  ab  to  median  of  a. 
Female  pores  are  an  inconspicuous  pair  in  XIV,  anteromedian  of  a.  Spermathecal 
pores  on  small  papillae. 

Internal  anatomy  (paralectotype  i ;  gross  anatomy  confirmed  from  all  localities 
and  details  from  14/4,  specimen  12)  :  supra-oesophageal  vessel  in  £VIII— |XIII, 
moderately  developed.  Oesophagus  somewhat  dilated  in  VIII ;  in  IX-XIII 


TASMANIAN   EARTHWORMS  285 

moderately  and  uniformly  dilated  and  slightly  moniriform  ;  in  XIV  and  XV  more 
slender.  Intestinal  typhlosole  absent  though  in  specimen  12  a  low  diffuse  dorsal 
ridge  is  perceptible  from  XXIV  posteriorly. 

Nephridia  :  vesiculate  stomate  and  astomate  nephridia  and  several  pairs  of 
meronephric  buccal  tufts.  The  ovoid-subspherical  bladders  each  receive  a  single 
duct  from  a  simple  or  completely  coiled  nephridium.  The  arrangement  of  the 
bladders  is  as  follows  :  at  the  posterior  end  of  the  body  there  are  5  (presetal)  bladders 
on  each  side,  one  in  each  of  the  four  setal  rows  and  a  supernumerary  bladder  in  cd, 
giving  10  bladders  per  segment ;  the  nephridia  supplying  these  posterior  bladders 
are  small  and  compact  but  there  is  in  addition  an  enlarged  nephridium  (megamero- 
nephridium),  with  a  preseptal  funnel,  the  duct  of  which  contributes  to  a  longitudinal 
excretory  duct  (ureter),  passes  through  many  caudal  segments  and  appears  to  dis- 
charge at  junction  of  rectum  and  body  wall.  In  the  forebody  a  bladder  persists  in 
each  of  the  four  setal  lines  of  a  side  but,  in  addition  1-3  supernumerary  bladders  are 
present  in  cd  and  I  or  2,  usually  2,  between  d  line  and  the  middorsal  line  ;  very 
occasionally  there  is  a  single  supernumerary  bladder  in  be.  The  number  of  bladders 
on  each  side  in  the  forebody  thus  varies  from  4  to  10.  No  bladders  are  detectable  in 
segments  I  and  II  (paralectotype  i)  but  one  is  present  in  d  in  specimen  12,  and  the  a 
bladder  appears  to  be  absent  in  III  and  is  not  observable  in  paralectotype  i,  though 
present  in  specimen  12  in  the  anterior  spermathecal  segment,  VIII.  Where  two 
bladders  are  present  above  d,  one  is  anterior  in  the  segment  and  the  other  is  equatorial 
and  less  lateral.  Sporadically  in  paralectotype,  but  not  in  specimen  12,  there  are 
two  bladders  in  c,  one  anterior  and  one  shortly  presetal.  Numbers  of  bladders 
counted  on  one  side  in  paralectotype  i  are  :  I,  o  ;  II,  o  ;  III,  4  ;  IV,  6  ;  V,  9  ; 
VI,  9  ;  VII,  8  ;  VIII,  7  ;  IX,  9  ;  X,  10  ;  XI,  10  ;  XII,  10  ;  XIII,  9  ;  post-pros- 
tatic,  8  ;  caudal,  5  ;  these  numbers  agreeing  well  though  not  always  exactly  with 
those  in  specimen  12.  Large  paired  tufted  nephridia,  each  consisting  of  numerous 
spiral  loops,  lie  in  segments  II-IX  ;  each  tuft  in  III-IX  contributes  to  a  composite 
duct  which  is  continuous,  on  each  side  of  the  nerve  cord,  from  segment  IX  to  the 
lower  lip  of  the  mouth  (paralectotype  i)  or  the  buccal  cavity  (specimen  12).  Each 
of  the  pair  of  tufts  in  II  sends  a  separate  composite  duct  to  the  lower  lip,  lateral  of  the 
other  longitudinal  duct  and  sends  a  few  very  slender  ducts  laterally  to  the  body  wall 
presetally  in  c  although  no  bladder  is  detectable. 

Simple  large  saccular  seminal  vesicles  in  IX  and  XII.  Prostates  irregular,  lobu- 
lated  to  racemose,  linear  structures,  extending  through  XVIII-XXI  (XVIII-XIX  in 
paralectotype  3)  ;  each  with  a  sinuous  narrow  muscular  duct  which  is  joined  entally 
near  its  junction  with  the  gland,  by  the  vas  deferens.  In  transverse  section  (paralec- 
totype i)  each  gland  is  seen  to  have  a  narrow  central  lumen  around  which  are  many 
distinct  groups  of  glandular  cells,  each  group  with  the  appearance  of  a  liver  lobule. 
The  cells  of  the  lobule  radiate  from  a  central  axis  which  in  some  cases  can  be  seen  to 
form  a  definite  lumen  although  this  is  not  lined  by  a  recognizable  epithelium.  The 
epithelium  of  the  central  lumen  of  the  gland  is  sporadically  interrupted  to  receive 
the  lumina  or  axial  cores  of  the  lobuli.  Width  of  the  gland  :  width  of  the  central 
lumen  =  1-61  :  0-18  mm  =  9:1.  Penial  setae  stout,  curved,  tapering  ectally  to  a 
simple  blunt  tip  ;  the  ectal  tapered  region  strongly  convex  '  dorsally ',  less  convex,  to 


286  B.    G.   M.    JAMIESON 

concave,  'ventrally'.  The  ectal  region  typically  (paralectotype  3)  ornamented  with 
approximately  16  circlets  spaced  at  regular  intervals  averaging  10  /zm,  each  circlet 
serrated  by  very  numerous  simple  minute  contiguous  spines  ;  the  spines  anteriorly 
directed,  adpressed  to  the  surface  of  the  seta,  only  slightly  notching  the  seta  in  profile. 
Specimens  from  other  localities  are  similar  in  general  form  and  may  have  similar 
ornamentation,  or  the  circlets  may  be  interrupted  in  such  a  way  that  the  different 
portions  are  not  always  in  the  same  transverse  line,  and  in  some  specimens  (e.g.  7/2, 
specimen  i)  there  is  no  regular  transverse  arrangement  of  spines  but  merely  scattered 
short  transverse  groups.  Sometimes  (13/4,  specimen  5)  there  are  short  spine-like 
ridges  adherent  longitudinally  to  the  setal  surface  immediately  ectal  to  the  spines 
(and  also  a  tendency  to  longitudinal  furrowing  of  the  seta)  ;  the  presence  of  ridges  in 
this  specimen  approaches  the  condition  characteristic  of  C.  polynephricus  poly- 
nephricus  ad  urethrae  (q.v.)  ;  lengths  of  well  developed  setae  073-1-5  mm,  general 
width  of  shaft  23-37  /um  (paralectotype  3  ;  specimens  I,  5,  10,  14,  18,  24,  34,  42). 
Metagynous  ;  ovisacs  not  developed  (paralectotype  i)  or  present  (specimen  12  and 
other,  but  not  all,  specimens).  Spermathecal  diverticulum  (inseminated)  digitiform. 

TYPE-LOCALITY.     Mt  Wellington ;  Hobart.     Map,  14/4.      Parattah.     Map,  14/1. 
MATERIAL  EXAMINED. 

Mt  Wellington,  I47°I5'E.  42°55'S.  Map,  14/4,  NMV :  01436,  3  paralectotypes.*  Fern 
Glade,  Emu  R.,  Burnie,  under  logs,  I45°55'E.  4i°o5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  17  May 

1954  ~  specimens   1-4  BM(NH)  :  1973.2.11-14.     Tarraleah,   over  pipeline,    I46°25'E.  42°2o'S. 
Map  13/4,  Dr  J.  L.  Hickman,  27  May  1954  -  specimens  5-6  BM(NH)  :  1973.2.15-16.     Collins- 
vale,  near  Hobart,  myrtle  forest,  I47°I5'E.  42°5o'S.     Map,  14/3,  Dr  J.  L.  Hickman,  8-9  Nov 

1955  -  specimens  7-11  BJ  :  T.38-42.     Mt  Wellington,  I47°I5'E.  42°55'S.     Map,  14/4,  Dr  J.  L. 
Hickman,  27  Oct  1955  -  specimens  12  and  13  BJ  :  7.39-40  ;   under  stones  on  top  of  mountain, 
Dr  J.  L.  Hickman,  13  Nov  1954  -  specimen  14  BJ  :  741  ;  specimen  15  AM  :  W52I3  ;  specimens 
16  and  17  BM(NH)  :  1973.2.17-18.     Shoobridge  Bend  track,  altitude  approximately  580  m, 
in  loam  and  clay  in  eucalypt  -  fern  woodland,  Dr  B.  G.  M.  Jamieson  and  Mr  E.  A.  Bradbury, 
19   Aug   1971  -  specimens    18-23   BJ  :  742-47.     Hobart,   Waterworks   Road,   under  stones, 
i47°2o'E.  42°5o'S.     Map,    14/5,   Dr  J.   L.   Hickman,    17  Aug   1954  -  specimen  24  BM(NH)  : 
1973.2.19  ;    specimen  25  BJ  :  748.     Lenah  Valley,  near  Newton  Falls,  under  stones,  Dr  J.  L. 
Hickman,  24  Jun  1957  -  specimen  26  AM  :  W52I4  ;   specimens  27-33  AM  :  53o5-53ri  '•   Lenah 
Valley,  New  Town  Creek,  Dr  J.  L.  Hickman,  15  Sep  1953  -  specimens  34-36  TM  :  1^331-333  ; 
31  Aug  1953  -  specimen  37  TM  :  1^334.     'Tasmania',  Dr  J.  L.  Hickman,  specimens  38-41 
TM  :  K335-338. 

REMARKS.  In  the  above  account  the  occurrence  of  a  pair  of  longitudinal  excretory 
ducts  and  associated  stomate  megameronephridia  is  demonstrated  for  the  first  time. 

C.  polynephricus  polynephricus  is  distinguishable  from  C.  p.  urethrae  and  from  the 
closely  related  C.  enteronephricus  in  location  of  the  anterior  genital  markings,  or  at 
least  those  in  IX,  significantly  lateral  of  the  spermathecal  pores.  The  similarity, 
nevertheless,  of  the  genital  fields  and  general  anatomy  of  C.  polynephricus  and  C, 
enteronephricus,  and  the  intermediate  position  of  C.  p.  urethrae  (q.v.)  indicate  specia- 
tion  from  a  common  origin  within  the  genus. 

*  The  specimen  designated  the  lectotype  by  Jensz  and  Smith  (1969),  041,  is  here  shown  to  be  a 
specimen  of  Perionychella  (V  esiculodrilus)  mortoni.  This  specimen  is  clearly  not  a  syntype  as  it  did  not 
contribute  to  the  type-description  of  C.  polynephricus.  Smith  has  agreed  that  its  designation  as  the 
lectotype  was  invalid  according  to  Article  74  (a)  (i)  of  the  International  Code  of  Zoological  Nomenclature. 


TASMANIAN  EARTHWORMS 


287 


sp.p 


1mm 


g.m 


B 


FIG.  23.     Genital  fields.     Cryptodrilus  polynephricus  urethrae  :   A,  paratype  8  ;  B,  paratype  20. 


288  B.   G.  M.   JAMIESON 

Specimens  14-17,  from  the  summit  of  Mt  Wellington,  and  specimens  24  and  25, 
from  Hobart,  display  a  distinct  genital  field,  lacking  genital  markings  in  IX,  and 
having  those  in  X  and  XI  further  median  than  is  usual.  They  are  also  charac- 
terized by  a  longitudinal  reduplication  of  the  nephridial  bladders  in  c  lines  but  the 
latter  condition  occurs  sporadically,  in  some  segments,  in  the  paralectotypes  of 
which,  though  they  are  incompletely  mature,  one  has  rudiments  of  a  normal  field, 
with  genital  markings  in  IX  and  X  well  lateral  of  the  spermathecal  pores.  As  the 
nephridial  condition  of  the  atypical  specimens  is  echoed  in  the  types,  and  in  view  of 
the  great  variation  in  the  genital  fields  reported  for  another  species  of  Cryptodrilus 
(C  .  fastigatus  v.  Jamieson,  iQ72b)  erection  of  a  separate  taxon  for  specimens  14-17, 
24  and  25  does  not  seem  warranted. 

Cryptodrilus  polynephricus  urethrae  subsp.  n, 
Fig.  2,  2iC,  23,  246,  3iJ-M  ;  Plates  44-59.     Table  9 
1  =  330,  160  mm  ;  w  =  12,  6  mm  ;  s  =  175,  156. 

Prostomium  epilobous  |,  §  ;  dorsal  tongue  moderately  narrow,  closed  and  (in 
holotype)  weakly  canaliculate.  Paired  elliptical  accessory  genital  markings  with 
centres  in  a  and  with  wide,  whitish  glandular  borders,  in  intersegments  9/10  and  10/11 
(holotype,  paratype  i)  and  11/12  (paratype  i)  or  unilateral,  on  the  right,  in  holotype. 
Similar  markings  centred  in  a  in  17/18  ;  in  ab,  nearer  a,  in  18/19  and  median  to  a  in 
19/20.  Female  pore  in  XIV,  paired  anteromedian  of  a  in  holotype  ;  unpaired,  mid- 
ventral,  in  paratype  i.  Spermathecal  pores  very  slightly  lateral  of  a. 

TABLE  9 

Intersetal  distances  in  segment  XII  in  13  specimens*  of 
Cryptodrilus  polynephricus  urethrae 


holotype 

maximum 

minimum 

mean 

standard  deviation 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

4-0 

i-i 

2-4 

2-3 

14-0 

2-6 

3-o 

i-i 

4-0 

i-i 

2-9 

3'3 

14-0 

3-i 

3-0 

i-i 

1-6 

0-6 

1-6 

1-9 

2-7 

1-9 

1-6 

0-6 

2-5 

0-9 

2-1 

2'3 

6-7 

2-3 

2-1 

0-9 

0-8 

0-2 

0-4 

0-4 

3'4 

0-3 

0-4 

O-2 

standardized  (u  =  100) 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

13-02 

3-72 

7-91 

7'44 

45-58 

8-37 

9-77 

4-19 

16-14 

5-83 

14-29 

15-92 

45-58 

15-92 

14-29 

5-83 

9-30 

3-45 

7-91 

7'44 

20-11 

8-37 

9-09 

3-45 

12-61 

4-71 

10-76 

12-20 

3I-9I 

12-19 

10-88 

4-75 

i-75 

0-72 

1-81 

2-74 

7-66 

2-43 

i-53 

0-69 

2-68 

I-OO 

2-28 

2-59 

6-77 

2-59 

2-31 

I-OI 

holotype 

maximum 

minimum 

mean 

standard  deviation 

interval/afe 

*  H,  paratypes,  i,  7,  8,  9,  10,  12,  13,  16,  18,  20,  21  and  22. 


Supra-oesophageal  vessel  recognizable  in  |VIII,  IX-XIII,  XIV  well  developed. 
Oesophagus  vascularized  and  dilated  segmentally  in  VIII-XIII,  XIV  ;   intestinal 


TASMANIAN   EARTHWORMS  289 

typhlosole  absent  or  a  rudimentary,  very  low  ridge  present  from  approximately 
XXIV  (paratype  i  and  holotype  respectively).  Nephridia  :  5  pairs  of  large  tufts, 
each  with  multiple  ducts,  in  II -VI ;  the  ducts  of  those  of  II  discharging  at  the  junc- 
tion of  mouth  and  body  wall ;  those  of  III-VI  sending  composite  ducts  to  a  simple, 
though  still  composite,  common  longitudinal  duct  which  passes  forwards  to  the 
anterior  end  of  the  pharynx  (i.e.  enteronephric)  ;  sparse  exonephric  micromero- 
nephridia  present  on  the  body  wall  lateral  to  each  tuft.  Nephridia  in  VII  forming 
smaller  tufts  which  discharge  exonephrically  in  this  segment  (and  holotype,  in  VI  ?) 
and  appear  to  send  some  ducts  to  the  common  duct  of  the  anterior  tufts  ;  a  tendency 
to  ventral  tufting  continues  but  decreases  to  the  anterior  intestinal  region  where  there 
are  approximately  10  (paratype  i)  to  15  (holotype)  exonephric  micromeronephridia 
on  each  side  ;  the  nephridia  discharging  presetally  in  a,  b,  be,  c,  cd  and  above  d. 
The  median  nephridium  of  the  first  intestinal  segment  (XVI)  is  the  first  to  have  a 
preseptal  funnel  (lying  in  XV)  (demonstrated  in  paratype  i).  This  arrangement  of 
nephridia  persists  posteriorly  but  the  stomate  nephridium  now  enlarges  to  become  a 
megameronephridium,  the  duct  of  which  discharges  in  a  line  together  with  one  or 
two  ast ornate  micromeronephridia.  Caudalty  there  is  an  astomate  micromero- 
nephridium  in  each  of  setal  lines  a,  b,  c  and  d  and  4  above  d  on  each  side  (holotype, 
there  being  few  if  any  above  d,  but  additional  intersetal  nephridia  in  paratype  i)  ; 
the  megameronephridium  contributes  its  duct  to  a  slender  longitudinal  excretory 
duct  (ureter)  of  which  there  is  one  on  each  side  of  the  ventral  nerve  cord ;  the 
posterior  end  of  this  ureter  was  traced  into  the  extreme  caudal  segments  but  its 
termination  was  not  elucidated.  Nephridial  bladders  totally  absent. 

Racemose  seminal  vesicles  in  XI  and  XII  (holotype,  paratype  i)  with  a  unilateral 
rudiment  in  IX  in  holotype.  Prostates  depressed  tubuloracemose,  in  XVIII-XIX 
(holotype) -XX  (paratype  i),  irregular  in  width  and  lobulated,  with  very  narrow 
central  lumen.  Penial  setae  moderately  stout,  ectally  strongly  curved,  the  tip, 
which  may  or  may  not  be  reflexed  dorsally  or  bent  ventrally,  strongly  flattened  and 
almost  parallel  sided,  only  slightly  tapering  ectally,  and  sometimes  ventrally  some- 
what inrolled  ;  the  end  almost  straight  or  convex  and  simple  or  denticulate  ;  orna- 
mentation ental  to  the  depressed  apical  region  (holotype)  consisting  of  numerous 
transverse  rows,  spaced  at  approximately  6  /mi  intervals,  each  row  minutely  serrate, 
the  teeth  of  the  serrations  very  numerous,  closely  continguous,  in  a  single  series, 
minute  but  attenuated  in  the  axis  of  the  seta,  short  ridges  like  a  fringe  present  ectal 
to  each  tooth  row  ;  the  setal  surface  also  with  numerous  longitudinal  furrows.  In 
other  specimens  the  tooth  rows  may  be  broken  up  into  short  transverse  or  oblique 
series  of  several  to  many  teeth  and  these  groups  may  lie  in  the  same  transverse  or 
oblique  line  as  adjacent  groups  or  may  be  scattered  apparently  randomly,  length  = 
2-1-3-9  mm,  general  width  of  shaft  =  39-45  /im  (holotype  ;  paratypes  i,  6,  20). 

Spermathecal  diverticulum  simple,  uniloculate  but  with  a  basal  swelling. 

VARIATION. 

Genital  markings.  The  genital  field  in  this  subspecies  is  distinguished  from  that 
of  C.  p.  polynephricus  by  location  of  the  post-spermathecal,  preclitellar  accessory 
genital  markings  in  or  median  to  the  longitudinal  lines  of  the  spermathecal  pores  ; 


2QO  B.   G.   M.    JAMIESON 

the  latter  pores  are  usually  in  or  very  slightly  lateral  of  a  lines.  In  paratypes  18  and 
19  the  genital  markings  are  further  lateral  in  ab,  but  are  not  significantly  lateral  of  the 
spermathecal  pores  as  these  are  similarly  displaced.  It  can  be  seen  in  the  following 
table  that  the  genital  fields  in  the  holotype  and  paratype  I  are  representative  of  the 
most  common  conditions  in  the  remaining  paratypes.  Paratypes  2,  6  and  22  are 
shown  to  have  atypical  fields  but  at  present  there  appears  to  be  no  justification  for 
giving  them  separate  taxonomic  status. 

Variation  in  the  genital  markings  of  26  specimens  of 
Cryptodrilus  polynephricus  urethrae 

Total  number 
Genital  markings  Specimen  of  specimens 

Small  elliptical  paired  in  ab  to  median  of 

a  at  hind  margin  of  VII  P6,  22  2 

VIII  P2,  22  2 

IX  P2  i 

X  P2  i 

Paired  intersegmental  in  line 

with  or  median  to  the  spermathecal 

pores  in    9/10  H     Pi,  3,  4,  5,  10,  n,  16,  16 

17,  18,  19,  20,  21,  23, 

25,  26 
lo/n  H     Pi-5,  10,  n,  12,  13,  14,  21 

16,  17,  18,  19,  20-26 
11/12  H(R)     Pi-5,  8,  12-14,  18-21,  18 

23-26 

12/13  P4.  5.  8,  12  4 

16/17  Pg  i 

17/18  H     Pi-5,  10-14,  I6-i9,  21-26  21 

18/19  H     Pi-6,  8,  10-12,  14,  16-19,  22 

21-26 
19/20  H     Pi-6,  8,  10-13,  16-19,  21  17 

20/21  P8  I 

21/22  P8  I 

H  =  holotype ;   P  =  paratype. 

Other  variation.  Anatomical  variation  in  this  subspecies  in  some  systems  exceeds 
what  would  normally  be  expected  within  a  species.  Thus  though  a  typhlosole  is 
absent  or  rudimentary  in  the  holotype  and  paratype,  respectively,  there  is  a  definite 
laminar  typhlosole  in  paratypes  2,  10,  n  and  20  and  appreciable  typhlosoles  are 
present  in  a  considerable  proportion  of  the  remaining  type-specimens.  In  specimens 
from  the  same  sample  (e.g.  13/3,  paratypes  10-12)  there  may  be  variation  from  a  deep, 
to  a  moderate,  to  a  very  rudimentary  typhlosole.  Other  morphological  variation  does 
not  appear  to  correlate  with  this  typhlosole  variation  and  all  specimens  are  distingui- 
shed from  the  typhlosolate  C.  enteronephricus  in  lacking  segmentally  enteronephric 
nephridia  and  in  having  a  ureter.  Variation  in  the  distribution  of  seminal  vesicles 
also  occurs  from  IX  and  XII,  to  IX,  XI  and  XII,  to  XI  and  XII  and,  most  remark- 
ably, the  female  pore  is  paired  or  (paratypes  i,  3,  4,  8,  12,  26)  unpaired,  median. 


TASMANIAN   EARTHWORMS  291 

Variation  in  setal  ratios  is  considerable  (Table  9)  and  one  of  the  more  extreme 
variants  (paratype  9)  has  the  last  hearts  in  XII. 

REMARKS.  The  location  of  the  preclitellar  genital  markings  approximately  in 
line  with  or  median  to  the  spermathecal  pores,  the  absence  of  nephridial  bladders 
and  the  widened  ends  of  the  penial  setae  distinguish  this  subspecies  from  the 
nominate  subspecies. 

MATERIAL  EXAMINED. 

Arne  Road,  E.  side  of  Styx  River  bridge,  approx.  1000  ft  into  rainforest,  I46°35'E.  42°5o'S. 
Map,  13/8,  Mr  D.  Frankombe,  24  Sep  1958  -  holotype  TM  :  K34I  ;  paratype  2  TM  :  K/342. 
Arne  Valley,  10  miles  from  Greeveston,  i46°5o'E.  43°io'S.  Map,  18/3,  Dr  J.  L.  Hickman, 
ii  Nov  1955  -  paratype  i  BM(NH)  :  1973.2.24  ;  paratypes  3-5  TM  :  1^343-345  ;  paratypes 
22-25  TM  :  K346-349.  Florentine  Valley,  surface  soil  under  litter  in  E.  regnans  forest,  under 
stones,  i46°25'E.  42°35'S.  Map,  13/3,  Mr  M.  Gilbert,  7  Mar  1956  -  paratype  6  BM(NH)  : 
1973.2.25:  Florentine,  Mr  J.  M.  Gilbert,  21  Jul  1955  -  paratype  7  TM  :  K35O  ;  Florentine, 
rainforest  soil,  Mr  J.  M.  Gilbert,  4  Aug  1955  -  paratypes  8  and  9  AM  :  ^^5313-5314  ;  Florentine 
Valley,  under  pieces  of  wood  and  bark,  Mr  J.  M.  Gilbert,  14-19  ?  Aug  1955  -paratypes  10-12 
AM  :  W53i5~53i7.  Tarraleah,  i46°25"  E.  42°2o'S.  Map,  13/4,  Professor  V.  V.  Hickman, 
Dec  1953  -  Jan  1954  ~  paratypes  13  and  14  BJ  :  T/52-53.  Dee  Bridge,  Lyell  Highway,  i46°35'E. 
42°i5'S.  Map,  13/7,  Dr  J.  L.  Hickman,  24  May  1954  -paratype  J5  BJ  :  T54.  Maydena, 
i46°4o'E.  42°45'S.  Map,  13/9,  Dr  J.  L.  Hickman,  15  Oct  1960  -  paratypes  16  and  17  BM(NH)  : 
1973.2.26-27.  Russel  River,  Mr  C.  D.  King,  16  Feb  1939  -  paratypes  18  and  19  BJ  :  T/55-56. 
Kallista,  near  Junee,  i46°35'E.  42°45'S.  Map,  13/11,  Mrs  Rodway,  Tasmanian  Biological 
Survey  :  Dei3,  19  Apr  1938  —  paratype  20  BJ  :  T57.  Snowy  Mts,  4000  ft,  i46°4o'E.  43°oo'S. 
Map,  18/2,  Tasmanian  Biological  Survey:  J8,  Mr  C.  D.  King,  19  Jan  1939  -paratype  21 
TM  : 


REMARKS.  The  separate  subspecies,  C.  polynephricus  urethrae  (and  possibly  it 
should  be  a  species)  is  recognized  for  specimens  which  have  anterior  genital  markings 
in  or  median  to  the  spermathecal  lines  and  which  lack  bladders  while  possessing 
ureters.  It  appears  to  be  linked  to  the  nominate  subspecies  by  intermediate  forms 
which  are  described  below. 

Whether  polynephricus  and  urethrae  are  to  be  retained  as  subspecies  or  should  be 
regarded  as  sibling  species  cannot  be  unequivocally  decided  at  present  as  knowledge 
of  them  is  limited  to  their  morphology  and  inadequate  geographical  evidence.  The 
map  (Fig.  2)  shows  that  they  are  largely  allopatric.  Their  morphological  similarity, 
which  exceeds  that  between  most  species,  and  the  occurrence  of  morphological  inter- 
mediates where  their  distributions  overlap,  are  taken  for  the  time  being  to  indicate 
that  they  are  subspecies. 


Cryptodrilus  polynephricus  polynephricus  ad  urethrae 

Fig.  2,  2iB,  246,  31!  ;   Plates  32-43.     Table  10 

Some  specimens  (42-45,  47,  49),  referable  to  C.  polynephricus  polynephricus  from 
the  appearance  of  the  preclitellar  genital  field,  differ  from  paralectotypes  and  other 
specimens  described  above  in  having  the  nephridial  bladders  limited  to  the  hindbody 
where  they  are  accompanied,  as  usual,  by  a  ureter.  These  specimens  are  here  in- 
cluded in  the  nominate  subspecies  but  are  distinguished  as  Cryptodrilus  polynephricus 


292 


B.   G.   M.   JAMIESON 
1 


sp.p 


FIG 


1mm 

24.     Genital  fields.     A,  Cryptodrilus  polynephricus  polynephricus,  specimen 
B,  C.  p.  p.  ad  urethrae,  specimen  49.     C,  C.  p.  urethrae,  paratype  i. 


12. 


polynephricus  ad  urethrae.  With  these  are  included  two  specimens  (46  and  48)  in 
which  genital  fields  are  undeveloped  but  which  have  the  same  nephridial  condition. 
This  'polynephricus  ad  urethrae'  morph,  in  addition  to  general  morphological  re- 
semblance to  C.  p.  polynephricus,  further  resembles  the  latter  in  having  seminal 
vesicles  in  IX  and  XII  and  simple-pointed  penial  setae  although  in  specimen  44 
this  shows  signs  of  the  flattening  characteristics  of  the  penial  setae  of  C.  p.  urethrae. 
(The  latter  specimen  is  exceptional  in  having  last  hearts  in  XII.)  The  ornamentation 
of  some  penial  setae  also  seems  to  be  intermediate  between  that  in  the  two  subspecies. 
As  shown  in  the  map  (Fig.  2),  the  distribution  of  polynephricus  ad  urethrae  is  inter- 
mediate between,  though  partly  overlapping,  the  distributions  of  the  two  subspecies, 
on  present  evidence.  Details  of  the  penial  setae  follow  :  setae  stout,  curved,  taper- 
ing ectally  to  a  simple  blunt  or  flattened  chisel-like  tip  ;  where  simple  tipped,  the 
ectal  tapered  region  strongly  convex  dorsally,  less  convex,  to  concave,  ventrally. 
The  ectal  region  ornamented  with  numerous  transverse  or  oblique  circlets,  at  inter- 
vals of  approximately  10  /urn,  each  circlet  serrated  by  very  numerous  spines  which  in 
some  specimens  (e.g.  13/3,  specimen  42)  are  two  deep  and  are  immediately  preceded 


TASMANIAN   EARTHWORMS  293 

ectally  by  short  spine-like  ridges  adherent  longitudinally  to  the  setal  surface.  Spines 
anteriorly  directed,  adpressed  to  the  setal  surface,  only  slightly  notching  the  seta  in 
profile.  The  circlets  in  some  (e.g.  13/3,  specimen  44)  broken  up  and  a  regular  trans- 
verse arrangement  scarcely  apparent.  In  14/4,  specimen  48,  the  setal  surface  bears 
numerous  longitudinal  furrows.  In  specimen  44  the  spines  are  simple,  and  ridges 
and  significant  furrowing  are  lacking  but  the  tip  differs  from  the  typical  polynephricus 
condition  in  being  flattened  ;  1  =  0-55-1-75  mm,  general  width  of  shaft  =  19-30  /mi 
(specimens  42,  44,  48,  49). 

TABLE  10 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  polynephricus  ad  urethroe 


specimen  42  1-3 

specimen  44  i-o 

specimen  45  i-o 

specimen  47  1-5 

specimen  48  1-3 

specimen  49  2-0 


mm 

ab 

be 

cd 

dd 

dc 

cb 

ba 

0-6 

i-i 

1-6 

2-6 

1-8 

i-i 

0-6 

0-4 

I-O 

I-O 

1-6 

i-o 

i-o 

0-4 

o-4 
0-6 

0-9 

i-o 
1-8 

2-3 

I-O 

1-7 

i-o 

0-4 
0-6 

0-7 

i-i 

1*5 

2'5 

J"5 

I-I 

0-7 

0-8 

1-5 

2-3 

3-4 

2-3 

i«5 

0-8 

standardized  (u  =  100) 
ab  be  cd  dd  dc  cb  ba 


I2-OO 

5-33 

10-67 

15-33 

24-00 

16-67 

10-67 

5-33 

I4'I5 

5'37 

13-17 

I3-I7 

21-95 

13-66 

I3-I7 

5-37 

14-21 

5-58 

13-20 

14-72 

18-78 

14-72 

13-20 

5-58 

13-25 

5-36 

11-99 

16-09 

20-82 

I5'I4 

11-99 

5-36 

12-46 

6-40 

10-44 

14-81 

24-24 

14-81 

10-44 

6-40 

13-73 

5-39 

10-29 

15-69 

23-53 

15-69 

10-29 

5-39 

I3-30 

5-57 

11-63 

14-97 

22-22 

15-12 

11-63 

5-57 

2-39 

i-oo 

2-09 

2-69 

3-99 

2-71 

2-09 

I-OO 

specimen  42 
specimen  44 
specimen  45 
specimen  47 
specimen  48 
specimen  49 
mean 
interval/aft 

MATERIAL  EXAMINED. 

Florentine  Valley,  surface  soil  under  litter  in  E.  regnans  forest,  under  stones,  i46°25'E.  42°35'S. 
Map,  13/3,  Mr  M.  Gilbert,  7  Mar  1956  -  specimens  42  and  43  TM  :  1^339-340  ;  specimens  44  and 
45  BJ  :  T49-50.  Russell  River,  i46°4o'E.  42°45'S.  Map,  13/9,  Mr  C.  D.  King,  16  Feb  1939- 
specimen  46  BM(NH)  :  1973.2.20.  Mt  Wellington,  i47°i5'E.  42°55'S.  Map,  14/4,  Dr  J.  L. 
Hickman,  13  Jan  1954  -specimen  47  AM  :  W53I2  ;  Betts  Vale  at  creek,  in  moss,  under  logs 
and  stones,  Dr  J.  L.  Hickman,  4  Mar  1954  -  specimen  48  BJ  :  T$i.  Eagle  Hawk  Neck,  in  kelp 
on  rocks,  I47°55'E.  43°oo'S.  Map,  19/2,  Dr  J.  L.  Hickman,  Apr  1956  -  specimen  49  BJ  :  T5iA. 

Cryptodrilus  simsoni  Spencer,  1895 
Fig.  2,  25 A,  3iN,  O  ;  Table  n 

1  =  61  mm,  w  =  4,  4-1  mm,  5  =  95+  (posterior  amputee),  125  (lectotype, 
specimen  2). 

Prostomium  canaliculate,  epilobous  f ,  closed.  First  dorsal  pore  3/4.  Setae  8  per 
segment  in  longitudinal  rows,  d  irregular  posteriorly.  Nephropores  not  visible 


294 


B.   G.   M.   JAMIESON 


VIII 


B 


FIG.  25.     Genital  fields.     A,  Cryptodrilus  simsoni,  lectotype.   B,  Pinguidrilus  tasmanicus, 

lectotype. 

(specimen  2)  or  sporadically  visible  throughout  (lectotype)  and  especially  distinct  on 
and  shortly  behind  the  clitellum  where  in  each  segment  there  are  6  presetal  pores  on 
each  side  ;  i  median  to  a,  i  in  each  of  a,  b,  c  and  d  and  i  near  the  dorsal  midline 
(lectotype) .  In  some  Fern  Dene  specimens  there  are  as  many  as  3  presetal  pores  in 
the  vicinity  of  each  of  setae  a  and  b  on  the  clitellum.  Clitellum  annular,  XIII- 
XVIII,  weak  in  XIII  and  XVIII  but  well  developed  between.  Male  pores  on  dome- 
shaped  papillae  with  diameter  equalling  ab  and  central  pore  in  a,  on  XVIII.  Paired 
eye-like  genital  markings  with  centre  in  b  at  19/20  and  in  ab  at  20/21  ;  a  median 
transverse  genital  marking  occupying  bb  in  11/12  (specimen  2)  or  10/11  (lectotype) 
bearing  a  row  of  approximately  10  small  circular  markings  corresponding  with  the 
intersegmental  furrow.  Female  pores  a  pair  of  small  slits  just  median  and  slightly 
anterior  to  a  on  XIV.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9,  on  small  papillae 
in  a  (specimen  2)  or  slightly  lateral  of  a  (lectotype). 

Last  hearts  in  XIII  (latero-oesophageal)  ;  connectives  to  dorsal  and  supra- 
oesophageal  vessels  present.  Supra-oesophageal  in  IX-XIII  (specimen  2)  ; 
indistinct  in  lectotype.  Gizzard  small,  easily  compressible,  in  V.  Extramural 
calciferous  glands  absent.  Oesophagus  with  circumferential  vascular  striae  and 
internally  papilliform  rugae  in  X-XVI  (specimen  2),  the  rugae  becoming  definite 
lamellae  in  XIV-XV  and  less  distinct  in  XVI  in  lectotype.  Intestinal  origin  XVIII ; 
typhlosole  absent. 


TASMANIAN  EARTHWORMS 


295 


TABLE  n 

Intersetal  distances  in  segment  XII  in  Cryptodrilus  simsoni 

mm 


aa 

J'4 
i-i 
i-o 
i-i 
i-i 
i-o 


ab 
0-4 
0-6 


0-4 


be 
i-i 

1-2 
1-2 
I-O 
1-2 


cd 


i-i 


dd 

4-8 
4-7 
4'3 
4'4 
4'3 
3-8 

standardized  (u  =  100) 


dc 


i-i 


cb 
i-i 

1-2 
1-2 
I-I 
1-2 


ba 
0-4 

0-4 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

11-48 

3-36 

10-08 

II-2O 

38-64 

10-64 

10-08 

3-o8 

9-73 

5-oi 

9-37 

11-21 

39-23 

11-50 

9-44 

4-13 

9-15 

4-25 

n-ii 

10-13 

39-54 

10-46 

ii-ii 

4-25 

9-38 

3'75 

10-63 

11-56 

38-75 

11-56 

10-94 

3'44 

9-87 

4-14 

9-24 

11-78 

39-17 

11-78 

9-87 

4-14 

9-00 

4-82 

10-93 

12-86 

34-4I 

12-54 

10-61 

4-82 

9'77 

4-22 

10-23 

11-46 

38-29 

11-41 

io-34 

3-98 

2-32 

I-OO 

2-42 

2-72 

9-07 

2-70 

2-45 

0-94 

lectotype 
specimen  i 
specimen  2 
specimen  3 
specimen  4 
specimen  5 


lectotype 
specimen  i 
specimen  2 
specimen  3 
specimen  4 
specimen  5 
mean 
interval/aft 


Nephridia  :  in  the  forebody  and  anterior  intestinal  region  meronephridia  dis- 
charge presetally  in  each  of  a,  b,  c  and  d  lines  and  between  d  line  and  the  middorsum. 
Those  discharging  in  c  and  d  lie  far  below  these  loci  and  have  very  long  ducts.  It  was 
not  possible  to  demonstrate  unequivocally  that  there  were  two  nephridia  at  a  as  the 
external  pores  indicate.  In  the  most  anterior  segment  the  ventral  nephridia  form 
small  (exonephric  ?)  tufts  and  do  not  appear  to  be  accompanied  by  the  full  com- 
plement of  more  dorsal  nephridia  ;  in  the  intestinal  region  there  is  a  preseptal  funnel 
on  each  side  near  the  nerve  cord,  this  condition  persisting  to  the  caudal  extremity  ; 
median  nephridia  are  not  enlarged  relative  to  others  though  they  are  moderately 
large.  In  the  posterior  region  the  dorsalmost  nephridia  discharge  in  an  approximately 
straight  line  above,  in  or  below  d  lines  which  are  there  irregular.  In  the  midbody  a 
small  bladder-like  dilatation  (parasitic  artefact  ?)  of  a  nephridial  duct  is  sporadically 
present  though  most  ducts  do  not  reveal  even  an  intramural  bladder. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  elongate  racemose 
seminal  vesicles  in  XI  and  XII,  on  the  anterior  septa  ;  similar  structures  of  unknown 
function  are  present  on  the  anterior  septum  of  X  (specimen  2)  and  of  preceding  seg- 
ments, to  at  least  V  (lectotype) .  Metagynous  ;  ovisacs  absent.  Prostates  restricted 
to  XVIII,  tongue  shaped  racemose  but  with  a  short  narrower  ectal  portion  prior  to 
the  duct  and,  in  serial  sections,  revealing  a  single  narrow  central  lumen  with  cuboidal 
epithelium.  Vas  deferens  joining  the  ental  end  of  the  sinuous  duct.  Penial  setae 
moderately  slender,  curved,  needle  like  ;  the  tip  tapering  narrowly,  simply  but 
bluntly  pointed  ;  ornamentation  absent ;  length  =  1-2-1-4  mm  ;  general  width  = 
13-14  /am  (specimen  2).  Spermathecae  2  pairs.  Diverticulum  single,  clavate, 
uniloculate  (specimen  2)  or  circumscribed  by  a  furrow  so  that  it  has  a  slight  tendency 
to  be  bifid  (lectotype). 


296  B.    G.   M.    JAMIESON 

Genital  markings.  The  location  of  genital  markings  in  11/12  (in  contrast  with  10/11 
in  the  lectotype),  19/20  and  20/21  is  constant  in  the  9  Fern  Dene  specimens  ;  the 
markings  in  20/21  are  always  more  median  than  those  in  19/20. 

TYPE-LOCALITY.     Emu  Bay.     Map,  7/2.     Launceston.     Map,  9/5. 

MATERIAL  EXAMINED. 

Launceston,  i47°io'E.  4i°25'S.  Map,  9/5,  A.  Simson,  Feb  1892,  NMV  :  Gi 82 -lectotype. 
Fern  Dene,  Ironcliflf  Road,  near  Penguin,  I46°O5'E.  4i°o5'S.  Map,  8/1,  Collector  ?  13  Oct 
1954  -  specimens  1-3  TM  :  1^352-354  ;  4  BJ  :  T58  ;  specimens  5  and  8  BM(NH)  :  1973.2.29- 
30  ;  specimens  6  and  7  AM  :  W53i8-53i9  ;  specimen  9  BJ  :  T59. 

REMARKS.  The  transversely  papillate  or  punctate  ventral  genital  marking  in 
10/11  or  11/12  is  distinctive  of  this  species.  The  presence  of  dorsal  pores  also  dis- 
tinguishes it  from  all  Tasmanian  species. 

Genus  PSEUDOCRYPTODRILUS  Jamieson,  1972!} 

DIAGNOSIS.  A  pair  of  combined  male  and  prostatic  pores  on  XVIII.  Setae  8  per 
segment.  Nephridia  2  to  3  on  each  side  excepting  caudally  where  there  is  a  single  pair 
of  stomate  holonephridia  in  each  segment.  Typhlosole  weak  or  absent.  Prostates 
depressed  tubular. 

DESCRIPTION.  Terrestrial.  Dorsal  pores  present.  Setae  commencing  on  II, 
in  8  longitudinal  rows  which  may  become  irregular  posteriorly.  Ventral  setal 
couples  widely  paired  (aa  <  zab)  ;  setae  of  the  dorsal  couples  widely  separated, 
cd  >  2ab  and  approximately  as  far  apart  as  the  two  setal  couples  of  a  side  (cd  =  0-8- 
i-ibc)  ;  dorsal  median  intersetal  distance  approximately  one-third  of  the  circum- 
ference (dd  =  o-3«).  Setae  a  and  b  of  XVIII  replaced  by  penial  setae.  Nephro- 
pores  inconspicuous.  A  pair  of  combined  male  and  prostatic  pores  on  XVIII ; 
postclitellar.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9  or  anteriorly  in  VIII  and  IX. 

Dorsal  blood  vessel  single,  continuous  onto  the  pharynx  ;  last  hearts  in  XII  (latero- 
oesophageal).  Subneural  absent.  Gizzard  moderate  to  well  developed,  in  V. 
Calciferous  glands  absent  but  oesophagus  modified  (dilated  or  internally  rugose)  in 
XV-XVI,  XVII ;  intestinal  origin  XVIII  or  XIX  ;  a  low  typhlosole  present  or 
absent.  Anterior  nephridia  tuft-like  or  simple.  Nephridia  exonephric ;  in  the 
anterior  intestinal  region  2  on  each  side  ;  in  the  forebody  2  (P.  diaphanus)  or  3  (P. 
acanthodriloides,  in  which  the  median  one  is  stomate)  ;  caudally  holonephric,  with 
a  single  stomate,  avesiculate  nephridium  on  each  side.  Holandric  ;  gymnorchous  ; 
seminal  vesicles  in  IX  and  XII.  Prostates  depressed  tubular.  Penial  setae  present. 
Metagynous  ;  ovisacs  present.  Spermathecae  with  i  or  2  clavate,  uniloculate 
diverticula. 

DISTRIBUTION.  Eastern  Subregion  of  Australia,  southern  faunal  province : 
Victoria.  Tasmania.  (2  species.) 

TYPE-SPECIES.     Megascolides  diaphanus  Spencer,  1900. 

TASMANIAN  SPECIES.  P.  acanthodriloides  sp.  n. 

REMARKS.  The  new  species  is  included  in  the  formerly  monotypic  Pseudocrypto- 
drilus  chiefly  because  it  is  a  megascolecine  which  is  meronephric  anteriorly  but 


TASMANIAN  EARTHWORMS 


297 


1mm 


A 


sp.p 


1mm 


FIG.  26.     Genital  fields.     A  &  B,  Pseudocryptodrilus  acanthodriloides,  holotype  :   A,  entire 
male  field  ;   B,  detail  of  same.     C,  Rhododrilus  Httoralis,  holotype. 


18 


298  B.   G.  M.   JAMIESON 

holonephric  caudally.  This  condition  has  presumably  evolved  from  a  fully  holo- 
nephric  condition  and  may  have  been  convergently  acquired  in  the  two  species  but 
the  genus  as  redefined  above  to  include  acanthodriloides  is  satisfactorily  homo- 
geneous, there  being  no  variation  which  is  not  acceptable  as  intrageneric. 

The  longitudinal  orientation  of  the  accessory  genital  markings,  constituting 
tubercula  pubertatis,  is  highly  unusual  in  the  Megascolecidae  and  it  is,  therefore, 
remarkable  that  another  Tasmanian  species  Oreoscolex  bidiverticulatus  (q.v.)  has 
tubercula  and  an  almost  identical  genital  field.  Either  tubercula,  so  rare  elsewhere, 
have  been  acquired  independently  in  the  two  species  or  a  special  relationship  exists 
between  the  two  (i.e.  common  descent  from  a  species  with  similar  tubercula).  A 
decision  between  the  two  alternatives  cannot  be  made,  at  least  on  present  evidence, 
but  the  morphology  of  the  Oreoscolex  species  is  not  inconsistent  with  relationship. 
Origin  of  Oreoscolex  (Megascolecini)  in  south-eastern  Australia  from  local  holonephric 
species  or  from  species  which,  like  P.  acanthodriloides,  were  partly  holonephric  and 
partly  meronephric  is  quite  conceivable.  Irrespective  of  its  relationships,  Pseudo- 
cryptodrilus  presents  a  most  interesting  transition  from  the  holonephric  to  the  mero- 
nephric condition. 

Pseudocryptodrilus  acanthodriloides  sp.  n. 

Fig.  4,  26A,  B,  328  ;  Plates  61-63.     Table  12 
1  =  70,  67  mm,  w  =  3-4,  3-2  mm,  s  =  127,  126. 

Prostomium  epitanylobous  with  grooved  peristomium.  First  dorsal  pore  4/5. 
Setae  8  per  segment,  in  regular  longitudinal  rows  throughout.  Nephropores 
sporadically  visible  in  a,  b  and  d  lines.  Clitellum  annular,  ^XIII-XVII  (holotype  ; 
not  developed  in  paratype  i).  Male  pores  on  XVIII  in  a,  A  pair  of  longitudinal 
ridges  (tubercula  pubertatis)  present,  each  occupying  ab  and  extending  from  just 
behind  each  male  pore  to  the  setal  arc  of  XIX  ;  the  pair  of  tubercula,  with  the  male 
pores,  lying  on  a  whitish  glandular  tumescence  ;  a  similar  additional  pair  of  tubercula 
extending  from  the  male  pores  to  the  setal  arc  of  XVII  in  some  specimens  (see  field 
variation).  Female  pores  a  pair  shortly  anteromedian  to  a  of  XIV.  Spermathecal 
pores  2  pairs,  at  the  anterior  margins  of  VIII  and  IX,  but  not  in  the  intersegments, 
each  on  a  hemispherical  papilla  ;  the  ventral  surface  of  segments  VII-IX  and 
posterior  VI  tumid. 

TABLE  12 

Intersetal  distances  in  segment  XII  in  Pseudocryptodrilus  acanthodriloides 

mm 
aa  ab  be  cd  dd  dc  cb  ba 

holotype 
paratype  i 
paratype  2 
paratype  3 
paratype  4 


0-9 

o-5 

0-9 

0-9 

2-2 

0-9 

0-9 

o-5 

0-9 

0-4 

0-8 

0-9 

1-8 

0-9 

0-8 

0-4 

0-9 
0-8 

0-4 
0-4 

0-8 
0-8 

0-8 

I-O 

1-7 
1-8 

0-8 
i-o 

0-8 
0-8 

0-4 
0-4 

0-9 

0-4 

1*0 

0-9 

2-O 

0-9 

i-o 

0-4 

aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

12-04 

6-02 

"•57 

12-04 

28-70 

12-04 

ii-57 

6-02 

12-56 

6-03 

12-06 

12-56 

26-13 

12-56 

12-06 

6-03 

I3-83 

6-38 

11-70 

12-23 

25-53 

12-23 

11-70 

6-38 

11-44 

5'97 

11-94 

I3-43 

25-87 

I3-43 

11-94 

5-97 

11-90 

5'7i 

12-86 

11-90 

27-14 

11-90 

12-86 

5-7i 

12-35 

6-02 

12-03 

12-43 

26-67 

12-43 

12-03 

6-02 

2-05 

I  -00 

2-OO 

2-06 

4-43 

2-06 

2-OO 

I-OO 

TASMANIAN   EARTHWORMS  299 

Table  12  (cont.) 

standardized  (u  =  100) 

aa  ab  be 

holotype 
paratype  I 
paratype  2 
paratype  3 
paratype  4 
mean 
interval/aft 

Last  hearts  in  XII  (latero-oesophageal) .  Supra-oesophageal  in  VII-XII,  weakly 
developed.  Gizzard  moderate,  muscular  but  easily  compressible  in  V.  Extramural 
calciferous  glands  absent,  longitudinal  rugae  well  developed,  though  not  forming 
laminae,  in  XV  and  XVI.  Intestinal  origin  |XVIII ;  typhlosole  absent. 

Nephridia  :  exonephric  much  coiled,  tuft-like  nephridia  in  II,  III  and  IV  discharg- 
ing by  simple  ducts  presetally  in  c  and  or  d  lines.  Further  posteriorly  simple  avesi- 
culate  nephridia  discharging  presetally  in  association  with  setal  lines,  apparently  not 
all  4  setal  lines  in  any  one  segment  (a,  b  and  d  in  the  holotype  in  which  the  median- 
most  nephridium  is  seen  to  have  a  preseptal  funnel).  In  the  anterior  intestinal 
region  2  nephridia  on  each  side,  discharging  presetally  in  b  and  d  lines.  Caudally 
with  a  pair  of  stomate,  avesiculate  holonephridia  only,  in  each  segment,  discharging 
presetally  in  d  lines  (holotype  and  paratype  i).  Holandric  (funnels  iridescent  in  X 
and  XI)  ;  gymnorchous  ;  seminal  vesicles  racemose,  in  IX  and  XII.  Metagynous  ; 
small  ovisacs  in  XIV.  Prostates  flattened,  tubular,  tortuous,  in  XVIII  and  XIX  ; 
vas  deferens  joining  junction  of  gland  and  duct.  Penial  setae  moderately  slender 
but  relatively  short,  only  slightly  curved,  the  ectal  extremity  flattened  like  the  blade 
of  an  oar;  ornamentation  a  few  deep  half-encircling  jagged  scars  ;  length  =  0-6  mm; 
general  width  of  shaft  =  10  /zm  (paratype  3) .  Spermathecae,  2  pairs  ;  diverticulum 
single,  clavate,  uniloculate. 

Genital  markings.  Of  13  paratypes,  3  (paratypes  i,  7,  12)  have  male  fields  as  the 
holotype,  with  genital  markings  restricted  to  a  pair  of  tubercula  extending  from 
|XVIII  into  XIX  ;  6  paratypes  (2,  3, 4,  5,  n  and  13)  have  these  tubercula  augmented 
solely  by  a  pair  (unilateral  only  in  paratypes  3,  n  (R)  and  13  (L))  extending  from 
approximately  |XVIII-|XVII  which  lie  in  the  same  tumid  area  as  the  posterior 
pair  ;  in  i  paratype  (8)  the  posterior  tubercula  are  present  but  there  is  also  a  mid- 
ventral  circular  presetal  glandular  patch  in  XVIII  and  a  similar  postsetal  patch  in 
XVII  ;  i  paratype  (10)  has  the  posterior  tubercula  and  only  the  midventral  presetal 
patch  in  XVIII  ;  while  i  (paratype  6)  has  this  condition,  and  also  the  anterior  paired 
tubercula,  and  i  (paratype  9)  is  similar  to  paratype  6  but  lacks  the  left  anterior  tuber- 
cula. Most  specimens  have  ventral  tumescence  in  VII-IX  and  some  in  VI  also. 

MATERIAL  EXAMINED. 

Great  Lake,  shore  under  decaying  gum  leaves,  I46°45'E.  4i°55'S.  Map,  8/2,  Dr  J.  L.  Hickman, 
26  May  1954  -  holotype  TM  :  K355  ;  paratypes  i,  n  and  13  BM(NH)  :  1973.2.31-33  ;  paratypes 
2-5  TM  :  W356-359  ;  paratypes  6,  8-10  BJ  :  T6o,  61-63  ;  paratypes  7  and  12  AM  :  W532O- 
5321. 

REMARKS.     The  male  genital  field  distinguishes  this  species  from  P.  diaphanus. 


300  B.   G.   M.   JAMIESON 

Tribe  DIGHOGASTRINI  Jamieson, 
Genus  HICKMANIELLA  gen.  nov. 

DIAGNOSIS.  Combined  male  and  prostatic  pores  a  pair  on  XVIII.  An  oeso- 
phageal  and  an  intestinal  gizzard  present.  Nephridia  few  exonephric  avesiculate 
meronephridia  per  segment ;  the  medium  nephridium  with  preseptal  funnel. 
Prostates  tubuloracemose.  Spermathecae  2  pairs. 

DESCRIPTION.  Terrestrial.  Form  circular  in  cross  section ;  anus  terminal. 
Dorsal  pores  present.  Setae  numerous  in  each  segment  (>  30  in  posterior  segments)  ; 
ventral  gap  large  ;  dorsal  gap  recognizable  anteriorly,  inappreciable  caudally 
(zz  =  zy).  Nephropores  not  externally  visible.  Clitellum  annular  ;  anterior  to  the 
male  porophores.  Combined  pores  of  the  vasa  deferentia  and  a  pair  of  tubulorace- 
mose prostates  one  pair  on  XVIII.  Female  pores  a  pair  on  XIV.  Spermathecal 
pores  2  pairs,  in  intersegments  7/8  and  8/9. 

Last  hearts  in  XII  (latero-oesophageal).  Subneural  vessel  absent.  Oesophageal 
gizzard  in  V.  Extramural  calciferous  glands  absent.  Intestine  commencing  in  the 
vicinity  of  the  prostate  glands  ;  with  a  very  thick-walled  intestinal  gizzard  preceded 
by  a  thin-walled  crop  ;  typhlosole  and  caeca  absent.  Few,  exonephric,  avesiculate 
meronephridia  per  segment,  the  median  nephridium  in  oesophageal  and  intestinal 
regions  with  preseptal  funnel ;  tufting  and  enteronephry  absent.  Holandric ; 
gymnorchous  ;  seminal  vesicles  in  IX  and  XII.  Metagynous ;  ovisacs  absent. 
Penial  setae  present.  Spermathecae  with  clavate  but  internally  multiloculate 
diverticulum. 

TYPE-SPECIES.     Hickmaniella  ,opisthogaster  sp.  n. 

DISTRIBUTION.     Northwest  Tasmania.     (Monotypic.) 

REMARKS.  Hickmaniella  is  almost  unique  in  the  known  Megascolecidae  in  possess- 
ing an  intestinal  gizzard.  (Pleionogaster  has  3  or  4  such  gizzards.)  This  feature  is 
characteristic  of  the  Oriental  family  Moniligastridae  (in  which,  however,  Gates  (1962) 
states  the  gizzards  are  actually  oesophageal  though  post-ovarian),  the  holarctic  family 
Lumbricidae  and  the  palaearctic,  neotropical  and  Ethiopian  subfamily  Alminae  of 
the  non-Australian  family  Glossoscolecidae.  The  Moniligastridae  (Order  Moniligas- 
trida)  differ  from  Megascolecids  in  many  significant  respects,  including  the  single- 
layered  clitellum,  large-yolked  eggs,  presence  of  intra-septal  testis  sacs,  location  of 
one  or  two  pairs  of  male  pores  in  the  next  segment  or  segments  respectively  and 
absence  of  an  anterior  gizzard.  The  Lumbricidae  are  distinguished  by  the  absence 
of  an  oesophageal  gizzard,  location  of  the  male  pores  in  or  in  front  of  segment  XV, 
location  of  the  clitellum  far  behind  these  pores,  the  exclusively  lumbricine  setae  and 
in  other  respects.  In  contrast,  the  Glossoscolecidae,  while  showing  their  closest  affini- 
ties with  the  Lumbricidae  (vide  Jamieson  197 id),  show  notable  resemblances  to  the 
Megascolecidae  which  may  indicate  that  they  are  more  closely  related  to  the  latter 
than  is  any  other  family  with  the  exception  of  the  Eudrilidae.  The  intestinal  gizzard 
of  Hickmaniella  constitutes  a  further  link  between  the  Megascolecidae  and  the  Glossos- 
colecidae. Like  many  other  common  features  of  the  two  families  the  intestinal  gizzard 
may  have  been  independently  acquired  in  each  but  sufficient  similarities  exist  to 


TASMANIAN   EARTHWORMS  301 

suggest  that  if  this  be  the  case  we  have  here  a  case  of  parallelism  in  evolution  stemming 
from  a  common,  though  remote,  ancestry  rather  than  convergence  in  relatively  unre- 
lated lineages  of  the  order  Haplotaxida.  These  similarities  include  the  following  (in 
addition  to  general  characteristics  of  the  non-Alluroidid  Lumbricina  of  Brinkhurst  and 
Jamieson  1971)  :  progression  from  the  lumbricine  to  the  perichaetine  arrangement  of 
setae  ;  presence  of  prostate-like  glands  in  some  Glossoscolecidae  which  resemble  the 
prostates  of  the  Megascolecidae  ;  frequent  presence  in  both  families  of  oesophageal 
gizzards  and  the  mutual  possession  of  intestinal  gizzards  ;  location  of  the  male 
pores  in  XVIII  in  some  Glossoscolecidae,  as  in  most  Megascolecidae  ;  location  of 
these  pores  on  the  clitellum  in  some  Megascolecidae  as  in  many  Glossoscolecidae  and 
finally  replication  of  the  nephridia,  in  Tritogenia,  as  in  many  Megascolecids.  While 
inclusion,  on  the  one  hand,  of  the  Glossoscolecidae  and  Lumbricidae  in  the  super- 
family  Lumbricoidea  and,  on  the  other  hand,  of  the  Megascolecidae  and  Eudrilidae 
in  the  superfamily  Megascolecidae  is  here  upheld,  the  above-mentioned  similarities 
between  the  parent  families  Megascolecidae  and  Glossoscolecidae  bring  the  two 
families  closer  together  and  endorse  recognition  of  the  suborder  Lumbricina  as  a 
natural  grouping.  The  Alluroididae  included  in  it  by  Brinkhurst  and  Jamieson 
(1971)  relate  the  suborder  to  other  Haplotaxid  suborders,  especially  the  Haplotaxina. 


Hickmaniella  opisthogaster  sp.  n. 

Fig.  4,  i8A,  326,  D  ;  Plates  64-66 
1  =  61,  59  mm,  w  =  5-5,  5-0  mm,  s  =  87,  82. 

Prostomium  canaliculate,  tanylobous.  First  dorsal  pore  4/5.  Setae  on  XII  32 
(holotype)  or  49  (paratype  i)  ;  caudally  31  to  38  per  segment ;  aa  regular  through- 
out (holotype)  or  becoming  irregular  posteriorly  (paratype  i)  ;  anteriorly  aa  =•  $ab  ; 
caudally  aa  =  $ab  ;  zz  varying  from  2-2-^zy,  anteriorly,  to  i-i-$zy  caudally. 
Nephropores  not  externally  visible.  Clitellum  annular,  XIII-|XVII.  Male  pores 
approximately  in  a  lines  of  XVIII,  each  median  to  a  penisetal  pore,  on  prominent 
dome-shaped  medianly  almost  contiguous  papillae  which  expand  the  segment  longi- 
tudinally. Accessory  genital  markings  paired,  medianly  conjoined  oval  glandular 
prominences  with  papillate  or  pore-like  centres  in  approximately  b  lines  in  inter- 
segments 11/12,  15/16  and  20/21,  resembling  the  male  porophores  in  appearance  ;  an 
additional  similar  marking  unilateral  on  the  left,  in  14/15,  and  on  the  right,  in  21/22 
in  paratype  i.  Female  pores  paired,  anterior  to  setae  a  on  XIV.  Spermathecal 
pores  2  inconspicuous  pairs,  in  7/8  and  8/9,  in  a  lines. 

Last  hearts  in  XII  (hearts  in  X-XII  latero-oesophageal,  with  connective  from 
dorsal  and  supra-oesophageal  vessel ;  supra-oesophageal  vessel  recognizable  in 
|VII-|XIV,  moderately  developed).  Oesophageal  gizzard  moderate,  in  V.  Extra- 
mural calciferous  glands  absent.  Intestine  commencing  in  XVIII  in  which  it  is 
thin-walled  and  crop-like  (holotype,  paratype  i)  ;  a  very  large,  extremely  thick- 
walled  intestinal  gizzard  in  XIX  (holotype)  or  XX  (paratype  i)  ;  typhlosole  absent. 
Few,  exonephric,  avesiculate  meronephridia  per  segment,  commencing  in  II ;  3  on 


302  B.   G.   M.   JAMIESON 

each  side  in  the  intestinal  region  ;  median  nephridium  with  preseptal  funnel  in 
intestinal  and  at  least  the  posterior  oesophageal  segments  ;  tufting  and  enteronephry 
absent. 

Holandric  ;  gymnorchous  ;  seminal  vesicles  in  IX  and  XII.  Metagynous  ; 
ovisacs  absent.  Prostates  tubuloracemose,  extending  laterally  in,  and  restricted  to, 
XVIII  ;  with  very  narrow  central  lumen  wihch  bears  narrower  epithelium-lined  side 
branches.  Penial  setae  (holotype)  2  mature  and  i  completely  developed  on  each 
side  ;  when  functional  1-6-17  mm  l°ng>  general  width  of  shaft  31  ^m  ;  gently  or 
strongly  curved,  widened  basally,  the  ectal  tip  flattened,  usually  curved,  and  with 
margins  slightly  inrolled  ;  the  ectal  region  of  the  shaft,  but  not  the  inrolled  region, 
ornamented  by  numerous  short,  transverse  minutely  serrated  linear  markings  of 
which  there  are  4  or  more  across  the  width  of  the  seta  ;  this  ornamentation  readily 
visible  under  the  higher  powers  of  the  light  microscope  without  recourse  to  oil 
immersion.  Spermathecae  2  pairs,  in  VIII  and  IX  ;  a  single  clavate  but  internally 
multiloculate  diverticulum  ;  the  duct  with  a  large  dorsal  swelling. 

Genital  markings.  The  four  mature  paratypes  (i,  3-5)  resemble  the  holotype  in 
having  paired  intersegmental  genital  markings  in  11/12,  15/16  and  20/21.  In 
addition  paratype  i  has  unilateral  genital  markings  in  14/15  (left)  and  21/22  (right) 
resembling  the  other  markings. 

MATERIAL  EXAMINED. 


Parrawe,  I45°35'E.  4i°2o'S.  Map,  7/3,  Dr  J.  L.  Hickman,  25  Aug  1954  -  holotype  TM  : 
paratype  i  BM(NH)  :  1973.2.34  ;  paratype  2  TM  :  K36i.  Table  Cape,  i45°45'E.  4i°oo'S. 
Map,  7/1,  Dr  J.  L.  Hickman,  24  Aug  1954  -  paratypes  3  and  4  AM  :  W5322-5323.  Hellyer 
Gorge,  i45°35'E.  4i°2o'S.  Map,  7/3,  Dr  J.  L.  Hickman,  28  May  1954  -  paratype  5  BJ  :  T64. 


Tribe  MEGASCOLEGINI  s.  Jamieson,  i97ia 
Genus  OREOSCOLEX  Jamieson,  ig73a 

DIAGNOSIS.  Setae  8  per  segment.  A  pair  of  combined  pores  of  vasa  deferentia 
and  racemose  or  tubuloracemose  prostates  on  XVIII.  Gizzard  in  V.  Extramural 
calciferous  glands  and  typhlosole  present  or  absent.  Meronephric  ;  anterior  neph- 
ridia  astomate,  avesiculate  and  exonephric,  or  enteronephric,  usually  tufted  ;  caudal 
nephridia  avesiculate,  exonephric  (or  enteronephric  ?),  with  numerous  preseptal 
funnels  on  each  side  in  each  segment.  Spermathecae  in  2  or  3  segments,  paired  or 
median  ;  diverticula  uniloculate  or  multiloculate. 

DESCRIPTION.  Terrestrial  worms.  Form  circular  in  cross  section  ;  anus  ter- 
minal. Prostomium  prolobous  to  tanylobous.  Dorsal  pores  present,  commencing 
in  3/4-20/21.  Setae  8  per  segment,  in  regular  longitudinal  rows  throughout  or 
becoming  irregular  posteriorly.  Setal  ratios  very  variable  aa  :  ab  :  be  :  cd  :  dd  = 
1-9-3-4  :  i  :  1-5-5-9  :  I'6-3'4  :  3-1-12-9  ;  dd:u  =  0-11-0-58  ;  cd  =  or  <  never  sig- 
nificantly >  be.  Clitellum  annular  or  less  commonly  saddle  shaped,  occupying  4-6^ 
segments,  its  posterior  limit  in  XVII-XIX.  Combined  pores  of  the  vasa  deferentia 
and  a  pair  of  prostates  in  XVIII ;  the  prostates  externally  racemose  or  lobulated 


TASMANIAN   EARTHWORMS  303 

tubular,  in  the  former  case  with  or  without,  in  the  latter  always  with  an  axial  lumen  ; 
this  lumen  possessing  side  branches.  Female  pores  a  pair,  anteromedian  of  setae  a, 
in  XIV.  Spermathecal  pores  unpaired  posteriorly  in  VII  and  VIII  (0.  imparicystis) 
or  paired  in  6/7,  7/8-8/9.  Last  hearts  in  XII  or  (0.  bidiverticulatus)  XIII ;  latero- 
oesophageal  hearts  (always  ?)  with  connectives  from  dorsal  and  supra-oesophageal 
vessels.  Subneural  vessel  absent.  Gizzard  large,  in  V.  Oesophagus  with  internal 
rugae,  or  laminae,  or  extramural  calciferous  glands,  intestine  commencing  in  XVI, 
XVII  or  XVIII ;  typhlosole  absent  (0.  peculiaris),  rudimentary  or  well  developed, 
commencing  i|-8  segments  behind  the  intestinal  origin.  Meronephric  ;  anterior 
nephridia  astomate,  avesiculate  and  exonephric,  or  enteronephric  (entering  pharynx 
and  buccal  cavity),  usually  tufted  ;  caudal  nephridia  avesiculate,  with  numerous 
preseptal  funnels  on  each  side  in  each  segment  and  exonephric  or  (0.  imparicystis) 
possibly  enteronephric.  Holandric  with  seminal  vesicles  in  IX  and  XII  or  rarely  in 
XI  and  XII  or  (0.  imparicystis,  leai)  XII  only ;  or  (peculiaris)  with  anterior  male 
organs  reduced  (incipiently  metandric)  gymnorchous  (testis-sacs  absent).  Metagy- 
nous  (ovaries  in  XIII)  ;  ovisacs  present  or  absent.  Vasa  deferentia  joining  the  mus- 
cular duct  of  the  prostate  at  midlength  or  entally,  or  the  ectal  extremity  of  the  gland. 
Penial  setae  present  or  (imparicystis)  absent.  Spermathecae  with  one  or  two  uni- 
loculate  or  multiloculate  diverticula. 

TYPE-SPECIES.     Oreoscolex  imparicystis  Jamieson  I973a.     New  South  Wales. 
DISTRIBUTION.     Eastern  Subregion  of  Australia  :  New  South  Wales  and  Tasmania. 


Tasmanian  species 

0.  bidiverticulatus  sp.  n. 

*0.  campestris  (Spencer,  1895) 

*0.  irregularis  (Spencer,  1895) 

*0.  leai  (Michaelsen,  1910) 

0.  longus  sp.  n. 

0.  peculiaris  sp.  n. 

0.  sexthecatus  sp.  n. 

*0.  wellingtonensis  (Spencer,  1895) 


Other  species 

Four  species  from  New  South  Wales  (examined  by  the  author  as  indicated)  in 
addition  to  the  type-species.  The  additional  species  accord  exactly  with  the  generic 
diagnosis  but  detailed  generic  characterization  is  not  attempted  here. 

*Notoscolex  grandis  Fletcher,  i887a  'Cotypes'  AM  :  W.I494- 
*Cryptodrilus  illawarrae  Fletcher,  iSSga  Mt  Kembla  AM  :  W.I3H. 
*Cryptodrilus  saccarius  Fletcher,  i887b  'Cotypes'  AM  :  W.I372. 
*Cryptodrilus  simulans  Fletcher,  i88gb  Syntypes  AM  :  W.I499- 

*  Comb.  nov. 


B.   G.  M.   JAMIESON 


XIX 


B 

.  km  t  " 

FIG.  27.     Genital  j&elds.     A,  Oreoscolex  bidiverticulatus,  holotype.     B,  O.  longus,  holotype. 


TASMANIAN  EARTHWORMS  305 

KEY  TO  TASMANIAN  SPECIES  OF  OREOSCOLEX 

Spermathecal  diverticula  uniloculate    .........  2 

Spermathecal  diverticula  multiloculate          ........  5 

Male  pores  in  ab  on  XVIII  ..........  3 

Male  pores  median  to  a  on  XVIII        .........  4 

Reniform  extramural  calciferous  glands  in  XV  and  XVI.  Fig.  27 A   O.  bidiverticulatus  sp.  n. 
Extramural  calciferous  glands  absent.     Fig.  276  .          .          .          .          .    O.  longus  sp.  n. 

Male  pores  almost  contiguous  medianly,  on  a  circular  porophore  median  to  b  lines. 

A  median  genital  marking  in  20/21.  Fig.  28C  .  .  O.  irregularis  (Spencer,  1895) 
Male  pores  shortly  median  to  a  lines  on  a  transverse  oval  porophore  which  extends 

lateral  of  b  lines  and  impinges  on  adjacent  segments.  Fig.  28B  .  O.  peculiar  is  sp.  n. 

Spermathecae  3  pairs.  Fig.  2gA O.  sexthecatus  sp.  n. 

Spermathecae  2  pairs  ...........  6 

A  pair  of  eye-like  genital  markings  present  in  ab  in  17/18  or  XVII  and  in  each  of 

several  intersegments  behind  XVIII.     Fig.  28A          .         O.  campestris  (Spencer,  1895) 
Eye-like  or  other  paired  genital  markings  absent  .......  7 

A  large  median  transverse  genital  marking  in  each  of  intersegments  15/16-18/19, 

extending  approximately  to  b  lines.     That  in   18/19  indistinct ;    that  in   17/18 

indistinct  posteriorly O.  leai  (Michaelsen,  1910) 

A  large  mid  ventral  glandular  pad  in  19/20  or  20/21  extending  laterally  beyond  b 

lines.     Fig.  296      ......          O.  wellingtonensis  (Spencer,  1895) 


Oreoscolex  bidiverticulatus  sp.  n. 

Fig.  3,  27A,  32  J  ;  Plate  67.     Table  13 
1  =  150,  106  mm,  w  =  4-9,  4-2  mm,  s  =  118,  141. 

Prostomium  epilobous  J,  open.  First  dorsal  pore  4/5.  Setae  8  per  segment  in 
regular  longitudinal  rows  throughout.  Nephropores  not  visible.  Clitellum  annular 
(£XIII  dorsally)  XI V-XVII ;  not  developed  in  paratype  i.  Male  pores  on  XVIII, 
in  ab,  on  long  tubercula  pubertatis  which  extend  from  the  setal  arc  of  XVII  to  that  of 
XIX  ;  a  glandular  depression  occupying  the  space  between  the  tubercula.  Female 
pores  paired  anteromedian  of  a  on  XIV.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9, 
on  small  papillae  in  a. 

TABLE  13 

Intersetal  distances  in  segment  XII  in  Oreoscolex  bidiverticulatus 

mm 
aa  ab  be  cd  dd  dc  cb  ba 

holotype  1-8  0-6  1-7  i-i  4-4  i-i  1-8  0-5 

paratype  i  1-5  0-5  1-4  i-o  4-0  i-o  1-4  0-5 

standardized  (u  =  100) 
aa  ab  be  cd  dd  dc  cb  ba 

holotype 
paratype  i 
mean 
interval/oft 


14-02 

4-58 

12-94 

8-63 

33'42 

8-63 

13-75 

4-04 

13-54 

4-31 

12-31 

8-92 

35-38 

8-92 

12-31 

4-31 

13.78 

4-45 

12-63 

8-78 

34-40 

8-78 

13-03 

4-18 

3-10 

I-OO 

2-84 

1-97 

7-73 

1-97 

2-93 

0-94 

B.   G.   M.   JAMIESON 


sp.p 


1mm 


FIG.  28.     Genital  fields.     A,  Oreoscolex  campestris,  specimen  i.     B,  O.  peculiaris,  holotype. 

C,  O.  irregular  is,  specimen  i. 


Last  hearts  in  XIII  (latero-oesophageal)  ;  supra-oesophageal  in  VII-|XIV, 
moderately  developed.  Gizzard  large  and  firm,  in  V,  with  anterior  rim.  A  pair  of 
large  reniform  extramural  calciferous  glands  in  each  of  segments  XV  and  XVI,  each 
joined  over  most  of  its  height  to  the  lateral  wall  of  the  oesophagus,  free  for  only  a 
short  extent  dorsally,  and  ventrally,  and  with  a  wide  communication  with  the 
central  lumen  of  the  oesophagus  ;  sessile,  lacking  a  duct.  Each  gland  with  numerous 
radial  longitudinal  laminae  which  span  the  lumen  of  the  free  part  of  the  gland. 


TASMANIAN  EARTHWORMS  307 

Intestinal  origin  XVIII  but  no  constricted  oesophageal  valve  at  17/18  ;  deep  dorsal 
typhlosole  commencing  in  XXIII,  continued  anteriorly  as  a  slight  ridge  to  XIX.  In 
II-VII  with  a  pair  of  exonephric  tufted  nephridia  with  numerous  anteriorly  running 
ducts  (always  ?)  opening  in  the  same  segment ;  the  tufts  largest  in  V  and  VI  and  very 
large  in  V.  Few  exonephric  meronephridia  on  each  side  in  succeeding  segments  until 
in  XV  and  XVI  there  are  very  deep  lateral  bands  of  exonephric  nephridia,  each  with 
several  tubules,  with  the  appearance  of  transverse  tufts  ;  meronephridia  again  few 
in  more  posterior  segments  ;  caudally  numerous  exonephric  meronephridia  on  each 
side  dependent  from  the  anterior  septum  and  with  many  preseptal  funnels  in  a 
transverse  row.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  seminal  vesicles  race- 
mose, in  IX  and  XII.  Metagynous ;  ovisacs  present.  Prostates  racemose ; 
restricted  to  an  enlarged  segment  XVIII  :  with  (paratype  2)  branched  internal  ducts 
and  lacking  a  central  lumen.  Penial  seta  almost  straight,  filiform,  tapering  ectally, 
lacking  ornamentation  ;  the  tip  broadened,  irregular  and  bent  relative  to  the  axis  ; 
length  left  seta  =  1*1  mm,  general  width  of  shaft  =  12  pm  (holotype).  Sperma- 
thecae  2  pairs  ;  each  with  I  (paratype  i)  or  2  (holotype)  (inseminated)  clavate, 
uniloculate  diverticula. 

MATERIAL  EXAMINED. 

Fern  Glade,  Emu  River,  Burnie,  i45°55'E.  4i°o5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  24  Aug 
1954  ~  holotype  TM  :  K362  ;  paratype  i  BM(NH)  :  1973.2.35  ;  paratype  2BJ  :  T65. 

REMARKS.  Intraspecific  variation  from  one  to  two  diverticula  on  a  spermatheca 
is  highly  unusual  in  the  Oligochaeta  but  there  is  no  doubt  that  the  holotype  and  para- 
types  are  conspecific.  Location  of  last  hearts  in  XIII  and  of  calciferous  glands  in  XV 
and  XVI,  together  with  the  configuration  of  the  genital  markings,  is  diagnostic  of 
this  species. 


Oreoscolex  campestris  (Spencer,  1895) 
Fig.  3,  28A,  32K  ;  Plates  68-72.     Table  14 

Cryptodrilus  campestris  Spencer,  1895  :  39,  PL  II,  fig.  13-15. 
Notoscolex  campestris  ;  Michaelsen,  1900  :  192  ;   Jamieson,  19710  :  78. 

1  =  87,  82  mm,  w  =  3-5  mm,  s  =  109,  114  (specimens  i  and  2).     (1  =  50-75  mm, 
w  =  3  mm,  Spencer.) 

TABLE  14 

Intersetal  distances  in  segment  XII  in  Oreoscolex  campestris 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

specimen  i  1-4  0-4  2-1  i-i  1-6  i-i  2-2  0-4 

specimen  2  1-5  0-5  2-5  i-o  i-i  i-o  2-5  0-4 

specimen  3  i-i  0-3  1-4  0-8  i-o  0-9  1-6  0-3 

specimen  4  1-6  0-5  2-4  i-i  1-6  1-2  2-3  0-5 

specimen  5  1-4  0-5  2-3  i-o  1-4  i-o  2-5  0-4 


13-79 

4-14 

20-00 

II-O3 

!5-i7 

10-34 

21-38 

4-14 

14-19 

4-73 

23-65 

9-46 

10-81 

9-46 

23-65 

4-05 

15-00 

4-00 

19-00 

11-00 

14-00 

I2-OO 

22-00 

4-00 

14-19 

4-52 

21-29 

9-68 

14-19 

10-97 

20-64 

4-52 

13-50 

4-73 

21-60 

9-45 

13-50 

9-45 

23-63 

4-°5 

14-13 

4-42 

21-11 

IO-I2 

13-53 

10-44 

22-26 

4-15 

3-20 

I-OO 

4-78 

2-29 

3-06 

2-36 

5-04 

0-94 

308  B.   G.   M.    JAMIESON 

Table  14  (cow/.) 

standardized  (u  =  100) 

aa  ab  be  cd  dd  dc  cb  ba 

specimen  i 
specimen  2 
specimen  3 
specimen  4 
specimen  5 
mean 
interval/aft 

Prostomium  indistinctly  tanylobous,  canaliculate  (epilobous  £,  Spencer).  First 
drosal  pore  4/5  (3/4  Spencer).  Setae  8  per  segment  in  regular  longitudinal  rows 
throughout,  dd  narrowing  caudally.  Nephropores  not  externally  visible.  Clitellum 
annular,  XIII-|XVIII  (XIII-XVII,  saddle  shaped  in  XVII,  Spencer).  Male  pores 
in  or  median  to  a  on  XVIII  on  a  pair  of  low  circular  porophores  ;  each  pore  preceded 
and  succeeded,  on  the  porophore,  by  the  pore  of  a  penisetal  follicle.  Paired  eye-like 
genital  markings  occupying  ab  at  17/18,  19/20,  20/21,  21/22  (specimens,  I,  2)  and 
22/23  (specimen  i).  (Anterior  XVII,  18/19  and  19/20,  Spencer.)  Female  pores, 
paired,  anteromedian  of  a  on  XIV.  Spermathecal  pores  two  pairs  on  hemispheroidal 
papillae  at  the  anterior  margins  of  VIII  and  IX,  just  lateral  of  a.  Last  hearts  in  XII 
(latero-oesophageal).  Supra-oesophageal  in  VIII-|XIII,  well  developed.  Gizzard 
firm  and  very  large,  in  V.  Extramural  calciferous  glands  absent ;  oesophagus 
internally  rugose  in  ?,  X-XV.  Intestinal  origin  £XVI  (XVI,  Spencer)  ;  a  very 
deep  j_-shaped  dorsal  typhlosole  commencing  in  XIX.  Nephridia  :  very  large 
pharyngeal  tufts  in  IV  and  V  ;  smaller  anterior  tufts  apparently  exonephric  ; 
caudally  with  a  transverse  row  of  numerous  preseptal  funnels  on  each  side,  the 
nephridia  exonephric.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ; 
seminal  vesicles  racemose,  in  IX  and  XII.  Metagynous  ;  ovisacs  present  in  XIV  (?). 

Genital  markings  in  18  specimens  of  Oreoscolex  campestris 
Genital  markings  Specimen  Total  number 

of  specimens 

16/17  J6  i 

17/18  1-18  18 

18/19  5,  13.  M  (L).  18  4 

19/20  1-18  1 8 

20/21  1-18  18 

21/22  1-3,  7-II,   12  (L),   14,   I6-I8  13 

22/23  I  I 

Prostates  restricted  to  an  enlarged  segment  XVIII  ;  broad  flattened  smooth  lobes, 
each  with  3  serial  portions  joined  together  by  narrow  alternating  bands  and  probably 
originating  from  a  tubular  type  ;  the  two  glands  completely  encircling  and  adpressed 
to  the  intestine  and  to  each  other  middorsally  ;  central  lumen  very  narrow,  with  few 
lateral  branches  ;  vas  deferens  joining  the  muscular  prostate  duct  at  its  junction 
with  the  gland.  Penial  setae  gently  curved,  moderately  stout,  slightly  tapering 
ectally,  the  tip  truncated  and  fluted  by  longitudinal  grooves  ;  the  ectal  region 


TASMANIAN   EARTHWORMS  309 

ornamented  with  the  exception  of  the  vicinity  of  the  tip  ;  ornamentation  consisting 
of  numerous  regularly  spaced  circlets  of  large  acute  flattened  spines  the  bases  of  which 
are  in  contact  with  adjacent  spines  in  the  circlet,  some  spines  bifid,  most  simple  : 
length  of  a  functional  seta  =  0-34  mm  ;  general  width  of  shaft  =  13  /mi  (2  specimens) . 
Spermathecae  2  pairs  ;  diverticulum  single,  sessile,  multiloculate. 

TYPE-LOCALITY.     Parattah.     Map,  14/1. 
MATERIAL  EXAMINED. 

Parattah,  under  moss,  in  earth  at  base  of  cliff,  also  in  earth  along  edges  of  logs,  i47°25'E. 
42°2o'S.  Map,  14/1,  Professor  V.  V.  Hickman  and  Dr  J.  L.  Hickman,  18  Aug  1954  ~  specimens 
i,  3-6  TM  :  1^363-367  ;  specimens  2,  7-11  BM(NH)  :  1973.2.36-41  ;  specimens  12-16  AM  : 
W5324-5328  ;  specimens  17  and  18  BJ  :  T66-&7. 

REMARKS.  The  syntypes,  in  the  National  Museum  of  Victoria,  048,  are  desic- 
cated, and  yield  no  significant  taxonomic  information. 

Oreoscolex  irregularis  (Spencer,  1895) 
Fig.  3,  28C,  32L.     Table  15 

Cryptodrilus  irregularis  Spencer,  1895  :  34~35.  PI-  I  ng-  I-3- 
Notoscolex  irregularis  ;  Michaelsen,  1900  :  191  ;   Jamieson,  19710  :  79. 

1  =  194,  195  mm,  w  =  9  mm,  s  =  120,  136  mm  (specimens  i  and  2  throughout 
this  account). 

Prostomium  epilobous  f  (ca.  \,  Spencer)  canaliculate,  open,  parallel-sided.  First 
dorsal  pore  4/5.  Setae  8  per  segment ;  all  rows  becoming  irregular  near  the  caudal 
extremity  (or  from  XIV,  Spencer)  ;  setae  a  and  b  absent  in  XVIII  although  their  loci 
are  lateral  of  the  male  pores.  Clitellum  annular  XIV-XVIII ;  some  clitellar  modi- 
fication dorsally  throughout  XIII ;  dorsal  pores  occluded,  setae  and  intersegmental 
furrows  retained  though  fainter  than  elsewhere.  Male  pores  minute,  almost  con- 
tiguous medianly,  in  a  circular  depression  (with,  in  specimen  i,  posterior  extension, 
i.e.  pyriform),  on  a  low  median  circular  porophore  which  almost  fills  segment  XVIII 
longitudinally  and  extends  to  a  lines  laterally.  A  transversely  oval  median  accessory 
genital  marking  with  crater-like  centre,  in  20/21,  extending  slightly  lateral  of  a  lines 
and  almost  reaching  the  setal  arcs  of  XX  and  XXI.  Female  pores  a  pair,  antero- 
median  to  setae  a  of  XIV.  Spermathecal  pores  not  demonstrable  externally  ;  from 
internal  dissection,  well  median  of  a  lines  if  not  medianly  united,  but  said  by  Spencer 
to  be  in  a  lines. 

TABLE  15 

Intersetal  distances  in  segment  XII  in  Oreoscolex  irregularis 

mm 
act  ab  be  cd  dd  dc  cb  ba 

specimen  i  2-2  i-i  1-9  1-7  14-4  1-7  1-9  I-i 

specimen  2  2-6  1-3  1-7  1.2  15-0  1-2  1-7  1-3 

specimen  3  2-6  1-3  2-1  2-6  15-7  2-6  2-0  1-3 

specimen  4  2-8  0-9  1-5  2-1  13-9  1-9  1-5  0-9 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

8-49 

4-n 

7-40 

5-58 

55-35 

6-58 

7-12 

4-11 

9-86 

4'93 

6-58 

4-66 

57-54 

4-66 

6-58 

4-93 

8-53 

4-27 

6-87 

8-53 

52-14 

8-53 

6-64 

4-27 

10-88 

3-63 

5-86 

8-37 

54HI 

7-25 

5-86 

3-63 

9'44 

4-24 

6-68 

7-04 

54-86 

6-76 

6-55 

4-24 

2-23 

I-OO 

1-58 

1-66 

12-94 

i-59 

J-54 

I  -00 

3io  B.   G.   M.   JAMIESON 

Table  15  (cow/.) 

standardized  (u  =  100) 
aa  ab  be 

specimen  i 
specimen  2 
specimen  3 
specimen  4 
mean 
interval/aft 

Dorsal  vessel  single,  continuous  onto  the  pharynx.  Last  hearts  in  XII  (hearts  in 
X-XII  latero-oesophageal,  with  dorsal  and  larger  supra-oesophageal  connectives)  ; 
supra-oesophageal  in  VIII-|XIII,  well  developed.  Gizzard  large  but  easily  com- 
pressible in  V  ;  oesophagus  dilated  and  vascularized  and  internally  rugose  in  IX-XV 
(-XIII,  Spencer),  but  calciferous  glands  absent.  Intestinal  origin  posterior  in  XVII 
(in  XVIII,  Spencer)  ;  typhlosole  rudimentary,  commencing  in  XXV  ;  muscular 
thickening  and  caeca  absent.  Nephridia  :  large  tufts  in  II-IX,  especially  large  in  II 
and  III ;  those  in  V-IX  exonephric  by  composite  ducts  in  their  own  segments  ;  those 
in  IV  sending  composite  ducts  forwards,  to  the  vicinity  of  the  pharynx  or,  II  and  III, 
of  the  buccal  cavity  ;  some  tufting  of  median  nephridia  persists  to  XIII  and  in  II- 
XIII  there  are  no  nephridia  lateral  to  the  median  nephridium.  In  XIV  the  nephridia 
begin  to  spread  laterally  to  form  a  parietal  band  of  astomate,  avesiculate,  exonephric 
micromeronephridia  and  by  XIV  these  bands  are  well  developed  and  so  dense  as  to 
have  the  appearance  of  transversely  expanded  tufts,  though  with  numerous  separate 
parietal  ducts  in  the  same  segment.  Caudally  with  transverse  bands  of  30-40 
avesiculate  exonephric  parietal  micromeronephridia  on  each  side,  each  with  a 
minute,  preseptal  funnel. 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles 
racemose  in  IX  and  XII  (XII  only,  Spencer).  Metagynous  ;  ovisacs  present. 
Prostates  each  a  broad  tongue-shaped  lobe,  with  surface  lobulations,  racemose  in 
appearance  but  internally  with  single  axial  duct  which  gives  off  lateral  branches  into 
the  gland  ;  double  vas  deferens  joining  the  middle  of  the  long  muscular  external  duct. 
Penial  setae  absent.  Spermathecae  2  pairs,  duct  about  as  long  as  and  clearly  de- 
marcated from  the  ampulla,  joined  near  its  ectal  end  within  the  body  wall  by  a 
single  clavate  (inseminated)  diverticulum  ;  the  duct  entering  the  body  wall  at  the 
lateral  border  of  the  ventral  nerve  cord,  near  the  midventral  line. 

Genital  markings.  Constant  though  not  always  equally  developed.  Crater  on 
male  porophore  is  pyriform  in  specimen  4  as  in  specimen  i. 

TYPE-LOCALITY.    Table  Cape.     (Types  in  NMV,  not  seen.) 

MATERIAL  EXAMINED. 

Table  Cape,  i45°45'E.  4i°oo'S.  Map,  7/1,  Dr  J.  L.  Hickman,  24  Aug  1954  -  specimens  I 
and  2  TM  :  1^368-369  ;  specimen  3  BJ  :  T68  ;  specimen  4  BM(NH)  :  1973.2.42. 

Oreoscolex  leai  (Michaelsen,  1910) 
Notoscolex  leai  Michaelsen,  1910  :  99-102,  Fig.  18,  19. 
1  =  310  mm,  w  =  7  mm,  s  =  152. 


TASMANIAN  EARTHWORMS  311 

Prostomium  prolobous  or  possibly  slightly  and  broadly  pro-epilobous.  Setae 
unusually  delicate,  in  the  mid-  and  hindbody,  with  the  exception  of  the  hind  fourth, 
regularly  arranged  and  widely  paired,  aa  :  ab  :  be  :  cd  :  dd  =  1-6:1:2:1:8.  Fur- 
ther anteriorly,  near  the  male  pores  each  ventral  pair  is  much  narrowed  but  the 
lateral  pair  is  not  (in  XIX  aa  =  $ab).  Clitellum  midventrally  insunk,  though  not 
saddle-shaped,  in  |XIII-XIX.  Male  pores  on  small,  almost  circular  papillae  on 
XVIII  in  ab,  about  one-twelfth  of  the  circumference  apart.  Large  unpaired,  mid- 
ventral  rounded  hexagonal  genital  markings,  extending  laterally  approximately  to 
setal  lines  b  and  leaving  only  a  small  region  of  the  segment  between  them  in  15/16, 
16/17,  17/18  and  18/19  ;  the  most  posterior,  in  18/19,  on^Y  indistinct,  that  in  17/18 
posteriorly  indistinctly  demarcated.  Female  pores  indicated  by  a  rather  long,  trans- 
verse furrow  anteriorly  in  XIV.  Spermathecal  pores  2  pairs,  in  7/8  and  8/9,  those  of  a 
pair  about  one-tenth  of  the  circumference  apart,  in  line  with  the  male  pores  (in  setal 
lines  ab  ?). 

Gizzard  large  in  V  ;  oesophagus  only  slightly  swollen  segmentally ;  calciferous 
glands  absent.  Intestine,  at  least  in  the  first  24  segments,  spiral.  Dorsal  vessel 
single.  Last  hearts  in  XII.  Meronephric  ;  a  large  number  of  nephridial  tubules  in 
each  segment ;  in  the  f orebody,  in  the  region '  of  thickened  septa,  nephridia  less 
numerous  and  aggregated  ventrally  to  give  a  spurious  appearance  of  holonephridia. 
Holandric  ;  2  pairs  of  sperm  funnels,  in  X  and  XI  ;  seminal  vesicles  racemose  in  XII. 
Prostates  slenderly  tongue-shaped,  long,  narrow  and  flattened ;  tubuloracemose, 
with  a  central  lumen  extending  almost  the  whole  length  of  the  glandular  part ;  the 
mass  of  the  glandular  part  composed  of  numerous  adpressed  parts  with  apparently 
epithelial  ducts  opening  into  the  main  duct.  Vasa  deferentia  of  a  side  entering  the 
internal  duct  of  the  prostate  separately  at  its  distal  fourth.  Penial  setae  approx. 
2  mm  long  and,  almost  throughout,  20  /um  wide  ;  gently  and  simply  curved,  ectally 
simply  and  slenderly  pointed.  Distinct  ornamentation  absent. 

Spermathecae  2  pairs,  duct  short,  conical,  bearing  a  multiloculate  diverticulum 
with  4  or  5  sperm  chambers. 

TYPE-LOCALITY.     Tasmania,  vicinity  of  Hobart. 
MATERIAL  EXAMINED. 
None  available. 

Oreoscolex  longus  sp.  n. 
Fig.  3,  276,  32N,  O  ;  Plates  73-80.     Table  16 
1  =  325,  215  mm,  w  =  6-0,  5-2  mm,  s  =  210,  231. 

Prostomium  epilobous  |,  open.  Dorsal  pores  indistinct ;  the  first  in  20/21. 
Setae  8  per  segment,  in  regular  longitudinal  rows  throughout.  Nephropores  not 
visible.  Clitellum  annular,  |XIII-XVIII,  weaker  ventrally  in  bb  (holotype),  not 
developed  in  paratype  i.  Male  pores  at  the  centres  of  prominent  transversely  con- 
joined papillae  occupying  ab  on  XVIII  ;  two  long  conspicuous  penial  setae  projecting 
in  the  vicinity  of  each  pore.  Accessory  genital  markings  :  large  oval  paired  pads  in 
16/17  and  19/20  extending  both  laterally  and  medianly  of  ab,  a  smaller  pair  in  9/10 


312  B.   G.   M.   JAMIESON 

in  ab,  and  a  median  ventral  pad  in  20/21  (holotype)  ;  or  median  pads  similarly 
extending  laterally  of  b  in  each  of  intersegments  12/13-16/17  and  paired  markings 
in  19/20  and  20/21  (paratype  i).  Female  pores  a  pair  anteromedian  of  a,  on  XIV. 
Spermathecal  pores  2  pairs,  in  7/8  and  8/9,  on  small  papillae  in  a. 

TABLE  16 

Intersetal  distances  in  segment  XII  in  Oreoscolex  longus 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

paratype  i  1-3  0-6  1-7  1-2  4-5  1-3  1-7  0-5 

paratype  2  1-4  0-6  1-9  1-2  5-4  1-4  1-8  0-6 

paratype  3  1-9  0-7  2-1  1-8  4-5  1-8  2-1  0-7 

paratype  4  1-7  0-6  2-1  1-4  5-0  1-4  1-6  0-6 

paratype  5  1-8  0-6  2-0  1-3  5-0  1-3  1-9  0-6 

standardized  (u  =  100) 

aa  ab  be 

paratype  i 
paratype  2 
paratype  3 
paratype  4 
paratype  5 
mean 
interval/aft 

Last  hearts  in  XII  (latero-oesophageal ;  connective  to  dorsal  vessel  not  recogniz- 
able). Supra-oesophageal  in  (?)-• |-XIII ;  well  developed.  Gizzard  in  V  but  septa 
5/6  and  6/7  very  delicate  and  with  more  posterior  septa  deflected  far  posteriorly  by  it  ; 
very  strong  and  firm,  fusiform  with  anterior  rim.  Extramural  calciferous  glands 
absent.  Oesophagus  especially  vascular  in  XI-XIII ;  in  IX-XIII  with  higher 
internal  lamellae  which  are  most  numerous  and  most  slender  in  XI  and  XII  in  which 
they  occlude  the  oesophageal  lumen  though  they  do  not  fuse  centrally.  Intestinal 
origin  XVII  (holotype),  or  XVIII  (paratype  i)  ;  a  deep  laminar  dorsally  typhlosole 
(at  first  rudimentary)  beginning  in  XXIV.  Nephridia  all  avesiculate  :  anterior 
nephridia  indistinct ;  no  enteronephry  detected.  From  approximately  XIV  pos- 
teriorly there  is  a  plumose  median  nephridium  on  each  side  which  has  a  single  pre- 
septal  funnel  but  which  discharges  exonephrically  by  several  ducts  which  converge 
to  the  body  wall  presetally  in  a  ;  other  astomate  meronephridia  discharge  presetally 
in  b,  c  and  in  d  or  above  d.  Caudally  there  are  several  (>  10)  preseptal  funnels  on 
each  side  in  each  segment,  located  in  ab  ;  the  meronephridia  being  exonephric 
(holotype). 

Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal  vesicles 
small,  racemose,  in  XI  and  XII  (holotype  ;  not  developed  in  paratype  i).  Metagy- 
nous  ;  ovisacs  present.  Prostates  very  tortuous,  slender,  lobulated  tubes  in  XVIII- 
XX  with  narrow  central  lumen.  Vasa  deferentia  of  a  side  running  together  then 
separating  before  joining  the  muscular  prostatic  duct  near  its  ental  limit.  Penial 
setae  curved,  moderately  stout,  slightly  tapering  ectally,  the  tip  flattened  and  widened 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

10-19 

4-41 

13-50 

9-37 

34-99 

10-19 

13-22 

4-13 

9-91 

4-44 

13-37 

8-39 

37-14 

9-91 

12-40 

4-44 

11-87 

4-57 

13-70 

11-42 

28-77 

11-42 

13-70 

4-57 

11-92 

3-97 

14-44 

9-97 

34-87 

9.97 

10-88 

3-97 

12-40 

3-95 

13-85 

8-93 

34<63 

8-93 

13-37 

3-95 

11-26 

4-27 

13-77 

9-62 

34-08 

10-08 

12-71 

4-21 

2-64 

I  -00 

3-23 

2-25 

7-99 

2-36 

2-98 

0-99 

TASMANIAN  EARTHWORMS  313 

to  form  a  blade  with  almost  straight  transverse  or  oblique  somewhat  crenulated  end  ; 
ectal  region,  with  the  exception  of  the  blade,  ornamented  ;  ornamentation  consisting 
of  numerous  regularly  spaced  circlets  of  large  acute  spines,  the  circlets  interrupted  so 
that  the  spines  are  in  groups,  of  approximately  5,  slightly  separated  from  other 
groups  in  the  same  circlet ;  some  spines  bifid,  most  simple  (holotype,  paratype  I,  7)  ; 
length  of  right  seta  =  1-6,  2-6  mm  (holotype,  paratype  i)  ;  general  width  of  shaft  = 
18,  25,  19  /zm  (holotype,  paratypes  i,  7).  Spermathecae  2  pairs  ;  diverticulum 
(inseminated)  digitiform  but  ectally  widening  greatly  to  form  a  bulbous  swelling  over 
the  pore  ;  no  distinct  duct  developed. 

Genital  markings  in  8  specimens  of  O.  longus 

Genital  markings 

Paired  in 

Paired  (often  conjoined)  in 


Total  number 

Specimen 

of  specimens 

9/io 

H 

P2-7 

7 

I2/I3 

Pi 

i 

13/14 

Pi 

i 

14/15 

Pi 

i 

15/16 

Pi 

i 

1  6/i  7 

H 

Pi-3,5-7 

7 

19/20 

H 

Pi  -7 

8 

20/21 

H 

Pi-3,P5-6,7(L) 

7 

21/22 

PS 

i 

Median  in 
H  =  holotype ;    P = paratype . 

MATERIAL  EXAMINED. 

Cox's  Bight,  I46°I5'E.  43°3o'S.  Map,  18/1,  Tasmanian  Biological  Survey,  Mr  C.  D.  King, 
Nov  1938  and  Jun  1939  -  holotype  TM  :  K37O  ;  paratypes  i  and  5  BM(NH)  :  1973.2.43-44  ; 
paratypes  2-4  TM  :  1^371-373  ;  paratype  6  AM  :  ^5329  ;  paratype  7  BJ  :  T6g. 

REMARKS.  O.  longus  is  identifiable  by  the  distinctive  genital  field  and  the 
bulbous  swelling  of  the  spermatheca  over  its  pore  with  an  accompanying  short, 
digitiform  diverticulum. 

Oreoscolex  peculiaris  sp.  n. 

Fig.  3,  286,  32M  ;  Plates  81-82.     Table  17 
1  =  46,  60  mm,  w  =  5-7,  6-0  mm,  s  =  146,  125. 

Prostomium  epilobous  \  with  suggestions  of  a  tanylobous  condition.  First  dorsal 
pore  (5/6  ?),  6/7.  Setae  8  per  segment,  b,  c  and  d  irregular  posteriorly  ;  ab  narrowing 
and  slightly  nearer  to  the  ventral  midline  as  XVIII  is  approached  from  anteriorly  or 
posteriorly,  but  absent  in  XVIII.  Nephropores  not  visible,  clitellum  annular 
XIII-XVII.  Male  pores  a  pair  in  XVIII  median  to  setal  lines  a,  at  the  centre  of  a 
large  oblong-oval  pad-like  glandular  protuberance  which,  basally  at  least,  includes 
the  setal  arcs  (and  setae  ab)  of  XVII  and  XIX  and  extends  laterally  almost  to  mid 
be,  4  circular,  translucent  pore-like  markings  present  on  the  pad  but  discernible  with 
difficulty,  two  at  the  anterior  and  two  at  the  posterior  limit  of  XVIII,  with  centres 
shortly  lateral  of  b  lines  (holotype,  paratype  i  ;  similar  in  paratypes  2-5  ;  less 

19 


i'3 

0-4 

3'7 

I-O 

3-o 

i-i 

3-4 

0-4 

1-6 

0-6 

4'3 

1-6 

2-5 

1-8 

4-2 

0-6 

0-8 

0-6 

2-1 

1-2 

1-9 

1-3 

2-1 

0-6 

0-7 

0-6 

3'3 

2-2 

1-8 

2-1 

3'4 

0-6 

aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

8-96 

2-99 

25-87 

6-97 

20-90 

7-96 

23-38 

2-99 

9-17 

3-33 

25-00 

9-17 

I4-58 

10-42 

24-58 

3*75 

7-48 

5-44 

19-72 

11-56 

17-68 

12-24 

20-40 

5'44 

4-87 

3-90 

22-40 

15-10 

12-18 

14-12 

23-38 

3-90 

7-62 

3-92 

23-25 

10-70 

16-34 

11-19 

22-94 

4-02 

1-94 

I-OO 

5-93 

2'73 

4-17 

2-85 

5-85 

1-03 

3i4  B.   G.   M.   JAMIESON 

developed  and  tending  to  be  divided  in  two  medianly  in  paratypes  6-8).  Female 
pores  a  pair  anterior  and  slightly  median  to  a  on  XIV.  Spermathecal  pores  difficult 
to  observe,  2  pairs  of  small  pores  in  a  at  7/8  and  8/9. 

TABLE  17 

Intersetal  distances  in  segment  XII  in  Oreoscolex  peculiaris 

mm 
aa  db  be  cd  dd  dc  cb  ba 

holotype 
paratype  i 
paratype  5 
paratype  6 

standardized  (u  =  100) 
aa  ab  be 

holotype 
paratype  i 
paratype  5 
paratype  6 
mean 
interval/aft 

Last  hearts  in  XII  (latero-oesophageal ;  connective  to  dorsal  vessel  a  delicate 
filament).  Supra-oesophageal  in  (?)-|XIV,  well  developed.  Gizzard  large  and 
firm,  with  anterior  rim,  in  V.  -Extramural  calciferous  glands  absent ;  vascularized 
with  high  rugae  in  VIII-XIV,  at  first  few ;  especially  numerous  in  XII-XIV  in 
which  they  form  flattened  papillae.  Intestinal  origin  XVII ;  spiral  from  XX 
posteriorly  ;  typhlosole  absent.  Nephridia  (holotype)  :  first  nephridia  large  entero- 
nephric  (pharyngeal)  tufts  in  V  ;  large  tufts,  decreasing  in  size  posteriorly,  in  VI- 
VIII,  those  in  VI  probably  enteronephric,  the  others  apparently  exonephric.  Small 
exonephric  tufts  in  IX-XII.  In  XIII  and  XIV  the  individual  tubules  and  ducts  of 
the  tufts  have  dissociated  so  that  the  meronephridia  begin  to  spread  laterally  and  by 
XV  numerous  discrete  micromeronephridia  form  bands  extending  far  laterally,  the 
nephridia  being  astomate,  avesiculate  and  exonephric  ;  approximately  15  micro- 
meronephridia on  each  side  in  each  segment  shortly  behind  the  prostates.  Abruptly, 
at  the  sixty-ninth  segment,  several  dorsal  nephridia  which  are  dependent  from  the 
anterior  septum,  become  enlarged  and  acquire,  each,  a  preseptal  funnel,  giving  several 
funnels  laterally  on  each  side  in  each  segment ;  a  single  preseptal  funnel  also  is 
present  on  the  medianmost  nephridium  (holotype  ;  caudal  arrangement  confirmed 
in  paratype  3).  Holandric  (but  incipiently  metandric  ?)  funnels  in  X  and  XI  but 
iridescent  in  XI  only  ;  gymnorchous  ;  seminal  vesicles  racemose,  in  IX  (holotype) 
and  XII  (holotype,  paratype  i).  Metagynous  ;  ovisacs  absent.  Prostates  tubulo- 
racemose,  running  laterally  in  and  restricted  to  XVIII ;  vas  deferens  joining  the 
gland  near  its  ectal  end.  Penial  setae  filiform,  the  tip  forming  a  small  rounded 
button,  ornamentation  absent  but  longitudinal  furrows  present ;  length  of  left  seta 


TASMANIAN   EARTHWORMS  315 

=  0-8  mm,  general  width  of  shaft  =  12  p.m  (holotype).     Spermathecae  2  pairs ; 
diverticulum  single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

Mt  Wellington,  Shoobridge  Bend  Track,  approx.  580  m,  in  loam  and  clay  in  eucalypt  -  fern 
woodland,  i47°i5'E.  42°55'S.  Map,  14/4,  Dr  B.  G.  M.  Jamieson  and  Mr  E.  A.  Bradbury, 
19  Aug  1971  -holotype  TM  :  K/374  ;  paratypes  i  and  3  BM(NH)  :  1973.2.45-46  ;  paratypes  2, 
6  and  7  TM  :  1^375-377  ;  paratype  4  AM  :  W5330  ;  paratype  5  BJ  :  T7O. 

REMARKS.  Location  of  the  male  pores  median  to  setal  lines  a  on  a  midventral, 
pad-like  glandular  protuberance  which  bears  pore-like  genital  markings,  diagnoses 
this  species.  The  specific  name  refers  to  the  restriction  of  multiple  caudal  nephro- 
stomes  to  a  dorsal  situation  distinct  from  the  ventral  funnel.  Other  peculiarities  are 
indicated  in  the  generic  definition. 


Oreoscolex  sexthecatus  sp.  n. 
Fig.  3,  2gA,  32?,  Q  ;   Plates  83-86.     Table  18 
1  ?  (posterior  amputee),  w  =  6-3  mm,  s  ?  (holotype). 

Prostomium  epilobous  f .  First  dorsal  pore  5/6.  Setae  8  per  segment ;  b,  c  and 
d  irregular  posteriorly.  Nephropores  not  visible.  Clitellum  not  developed.  Male 
pores  in  b  on  ovoid  papillae  which  extend  from  a  lateral  of  b.  Paired  eye-like  genital 
markings  with  centres  in  b,  radius  ab  or  less,  in  13/14,  14/15,  15/16,  16/17  and  19/20. 
Female  pores  paired  on  small  papillae  anterior  and  slightly  median  of  a  on  XIV. 
Spermathecal  pores  3  pairs,  in  6/7-8/9,  in  b  lines. 

TABLE  18 

Intersetal  distances  in  segment  XII  in  Oreoscolex  sexthecatus 

mm 

aa  ab  be  cd  dd  dc  cb  ba 

holotype  1-6  0-6  1-8  1-7  6-3  1-9  1-8  0-6 

standardized  (u  =  100) 
aa  ab  be  cd  dd  dc  cb  ba 

holotype  9-73          3-54        11-06        10-62        38-94        11-50        11-06          3-54 

interval/afc  2-75          i-oo          3-12          3-00        n-oo          3-25          3-12          i-oo 

Last  hearts  in  XII  (latero-oesophageal,  with  filamentous  connective  to  dorsal 
vessel).  Supra-oesophageal  in  ^VIII-|XIII,  well  developed.  Gizzard  large  and 
firm,  in  V  ;  preceded  in  IV  by  a  large,  thin-walled  proventriculus.  Extramural 
calciferous  glands  absent.  Oesophagus  vascular  in  VII-XVI,  with  internal  rugae, 
though  these  are  poorly  developed  in  VII-IX  and  nowhere  form  lamellae,  their 
greatest  development,  as  flattened  papillae,  being  in  XIV.  Intestinal  origin  XVII 
(?  macerated)  ;  typhlosole  ?  Nephridia  :  astomate,  avesiculate,  exonephric  mero- 
nephridia  in  II  to  the  amputation  at  about  XXV  ;  very  slender  ducts  entering  the 


B.   G.   M.   JAMIESON 


VII 


IX 


g.n 


FIG.  29. 


1mm 


Genital  fields.     A,  Oreoscolex  sexthecatus,  holotype.     B,  O.  wellingtonensis ,  14/5, 

specimen  i. 


sp.p 


body  wall  in  front  of  setal  follicles  a,  b,  c  and  d  and  near  middorsal :  the  a  and  b  and, 
anteriorly,  the  c  and  d  nephridia  forming  spiral-looped  aggregations  with  the  appear- 
ance of  tufts,  but  presetal  ducts  remaining  separate.  A  posterior  fragment  which 
corresponds  in  appearance  with  the  posterior  amputee  (there  being  only  one  other, 
very  small,  worm  in  the  sample)  has  5  preseptal  funnels  on  each  side  in  caudal  seg- 
ments, the  nephridia  remaining  exonephric.  Holandric  (funnels  and  free  sperm 
masses  but  no  iridescence  observed)  ;  gymnorchous  ;  very  large  racemose  seminal 
vesicles  in  IX  and  XII.  Metagynous ;  ovisacs  in  XIV.  Prostates  racemose, 
curved  linear,  extending  through  several  segments,  the  surface  deeply  lobulated ; 
the  central  lumen  moderately  wide  with  man}''  side  branches.  Penial  setae  almost 
straight,  moderately  slender,  scarcely  tapering  ectally  except  at  the  tip  which  is 
obtuse  and  rounded  (worn  ?)  ;  the  region  behind  the  tip  furrowed  ;  the  shaft  further 
entally  bearing  numerous  scattered  groups  of  small  but  attenuated  spines  which  are 
often  bifid  or  trifid  ;  length  =  1-5  mm,  general  width  of  shaft  =  18  /mi.  Sperma- 
thecae  3  pairs  of  unusual  form  :  duct  greatly  expanded  with  a  correspondingly  en- 
larged internal  lumen,  joined  by  a  bilobed  or  multiloculate  diverticulum  ;  the  sac-like 
ampulla  with  one  or  two  variably  developed  ectal  outpouchings. 


TASMANIAN   EARTHWORMS 


317 


MATERIAL  EXAMINED. 

Lake  St  Clair,  i46°io'E.  42°O5'S.  Map,  13/5,  Tasmanian  Biological  Survey  :  Jig,  Mr  R. 
W.  Kerr,  Feb  1941  -  holotype  TM  :  K^8. 

REMARKS.  The  combination  of  several  pairs  of  intersegmental  genital  markings, 
3  pairs  of  spermathecal  pores  and  ectal  outpouchings  of  the  spermathecal  ampulla 
distinct  from  spermathecal  diverticula,  diagnoses  this  species. 


Oreoscolex  wellingtonensis  (Spencer,  1895) 
Fig.  3,  296,  32R  ;   Plates  87-90.     Table  19 

Cryptodrilus  wellingtonensis  Spencer,  1895  :  43-44,  PL  3,  fig.  25-27. 
Notoscolex  wellingtonensis  ;   Michaelsen,  1900  :  192-193  ;   Jamieson,  19710  :  79. 

1  =  140,  155  mm,  w  =  5-8,  5-5  mm,  s  =  138,  149  (specimens  I  and  3). 

Prostomium  epilobous  \,  closed.  First  dorsal  pore  4/5,  3/4  ;  pores  very  conspicu- 
ous in  the  mid-  and  hind-body.  Setae  8  per  segment  in  regular  longitudinal  rows  ; 
d  occasionally  absent.  Nephropores  not  visible.  Clitellum  annular,  XIV-XVII. 
Male  pores  on  XVIII  in  a  on  a  large  glandular  pad  which  extends  laterally  beyond  b 
and  fills  the  segment  longitudinally  (specimen  i),  this  pad  replaced  in  specimens  3-5 
by  two  large  oval  papillae,  each  bearing  a  male  pore.  A  large  median  oblong-oval 
glandular  pad  in  19/20  (specimens  I,  2)  or  20/21  (specimens  3,  4)  extends  laterally 
beyond  b,  includes  the  setal  annulus  of  the  preceding  segment  and  reaches  that  of  the 
succeeding  segment  and  is  traversed  by  a  groove  corresponding  with  the  inter- 
segment. A  similar  pad  present  in  12/13  in  specimen  3  and  a  suggestion  of  a  small 
midventral  pad  median  to  a  in  17/18  in  specimens  1-3.  Female  pores  paired,  median 
to  and  only  slightly  anterior  to  a,  at  about  \aa,  on  XIV.  Spermathecal  pores  2  pairs 
on  small  papillae,  or  concealed,  in  7/8  and  8/9,  in  a  lines. 

TABLE  19 

Intersetal  distances  in  segment  XII  in  Oreoscolex  wellingtonensis 


specimen  2 
specimen  3 
specimen  4 


specimen  2 
specimen  3 
specimen  4 
mean 
interval/aft 


aa 
1-6 


ab 


0-6 


be 

2-1 

1-8 


cd 

2-6 
I'S 


<fti 

8-9 

4'1 
8-2 


dc 
i-i 

2-6 


cb 
1-4 

2-3 
1-8 


standardized  (u  =  100) 


ba 


0-6 


aa 

ab 

be 

cd 

dd 

dc 

cb 

ba 

9-79 

2-98 

8-51 

7-66 

52-77 

6-81 

8-51 

2-98 

7-96 

3-10 

13-27 

16-37 

25-66 

16-37 

14-16 

3-10 

11-16 

3-19 

9-96 

8-37 

45-82 

8-37 

9-96 

3-19 

9-64 

3-09 

10-58 

10-80 

41-42 

10-52 

10-88 

3-09 

3-12 

I-OO 

3-42 

3-50 

13-40 

3'4° 

3-52 

I-OO 

Last  hearts  in  XII  (latero-oesophageal) .     Supra-oesophageal  moderately  deve- 
loped, ending  at  f  XIII.     Gizzard  large,  firm  in  V  extending  posteriorly  to  the  level 


3i8  B.   G.  M.   JAMIESON 

of  XI,  oesophagus  virtually  suppressed  to  this  segment ;  in  XII-XIV  with  high 
vascular  internal  rugae,  intestinal  origin  XVI,  deep  dorsal  typhlosole  commencing  in 
XXIV  (specimens  I  and  3).  Nephridia :  pairs  of  very  large  tufted  nephridia  in 
II-V  with  composite  ducts  ;  those  of  II  apparently  opening  into  the  buccal  cavity  ; 
those  of  III  and  IV  and  possibly  V  running  forward  to  open  ventrolaterally  at  the 
mouth.  Nephridia  in  VI- IX  forming  smaller  tufts  exonephric  in  each  segment.  By 
XII  there  are  dense  lateral  bands  of  meronephridia ;  those  of  segments  X  and  XI 
being  intermediate  between  tufts  and  these  lateral  bands.  Caudally  with  very 
numerous  exonephric  (?)  micromeronephridia  on  each  side,  each  nephridium  with  a 
minute  preseptal  funnel  (specimen  i).  Holandric  (funnels  iridescent  in  X  and  XI)  ; 
gymnorchous ;  seminal  vesicles  racemose,  in  IX  and  XII.  Prostates  linear  but 
externally  racemose,  extending  laterally  in,  and  restricted  to,  XVIII ;  narrow 
central  lumen  present  with  few  side  branches.  Penial  setae  moderately  slender, 
sinuous,  the  tip  flattened  and  widened  to  form  a  blade  which  is  terminally  straight 
edged  with  a  few  irregular  teeth  ;  ectal  region,  with  the  exception  ot  the  blade,  with 
a  feathered  appearance  produced  by  a  series  of  numerous  composite  spines,  each  of 
which  is  approximately  triangular  but  has  several  long  and  often  branched  lateral 
pinnae  ;  length  of  left  seta  =  i-i  mm  ;  general  width  of  shaft  =  18  pm  (specimen 
i).  Metagynous ;  ovisacs  absent.  Spermathecae  2  pairs ;  diverticulum  (insemi- 
nated) sessile,  multiloculate. 

TYPE-LOCALITY.     Mount  Wellington.     Map,  14/4. 
MATERIAL  EXAMINED. 

Domain,  Hobart,  i47°2o'E.  42°5o'S.  Map,  14/5,  Dr  J.  L.  Hickman,  14  Aug  1954  ~  specimen  i 
TM  :  K379,  7  Nov  1955  -  specimen  2  AM  :  W533I.  Kelly's  Basin,  Port  Davey,  I45°55'E. 
43°2o'S.  Map,  17/1,  Tasmanian  Biological  Survey:  Mr  C.  D.  King,  Jan  1940  -  specimen  3 
BM(NH)  :  1973.2.47.  'Tasmania',  Dr  J.  L.  Hickman  -  specimen  4  BJ  :  Tji. 

REMARKS.  The  new  material  conforms  sufficiently  well  with  Spencer's  account 
for  there  to  be  little  doubt  of  conspecificity.  The  fact  that  one  of  the  new  specimens 
is  almost  topotypic  and  that  no  form  more  closely  resembling  Spencer's  description 
was  found  further  supports  the  identification.  Differences  in  Spencer's  account  are 
slight,  viz,  length  <  100  mm  ;  prostomium  only  slightly  epilobous  ;  male  and  sper- 
mathecal  pores  in  ab  ;  clitellum  extending  slightly  onto  the  dorsal  surface  of  XVIII. 
Accessory  genital  markings  are  said  by  Spencer  to  be  swollen  tumid  ridges  on  XVIII, 
XIX  and  XX,  marked  differences  from  the  present  specimens,  but  his  illustration 
(Plate  3,  fig.  25)  shows  a  single  intersegmental  marking  only,  in  19/20,  as  described  in 
this  account. 

Genus  MEGASCOLEX  Templeton,  1844 

DIAGNOSIS.  Setae,  at  least  in  the  mid-  and  hindbody  numerous  (more  than  8) 
per  segment.  Prostates  i  pair,  racemose,  their  pores  on  XVIII  or,  exceptionally,  an 
adjacent  segment ;  sometimes  associated  with  penial  setae.  Female  pores  paired  or 
exceptionally  unpaired.  Spermathecal  pores  usually  1-5  pairs  between  IV  and  IX 
(the  exceptions  are  the  few  cases  where  the  pores  are  fused  in  the  midline,  or  where 
they  are  numerous  on  each  side  in  each  segment  occupied).  One  gizzard,  in  V,  VI 


TASMANIAN   EARTHWORMS  319 

or  VII.     Calciferous  glands  present   or  absent.     Meronephric,   with  or  without 
enteronephric   nephridia.     Holandric,    rarely   metandric ;     testis-sacs   present   or 
absent.     Ovaries  in  XIII.     Spermathecae  with  or  without  free  diverticula. 
TYPE-SPECIES.    Megascolex  caeruleus  Templeton,  1844  (Ceylon). 

DISTRIBUTION.  Ceylon  and  India.  Australia,  with  Tasmania.  New  Caledonia. 
New  Zealand.  Norfolk  Island.  Annam. 

Tasmanian  species 
M.  burniensis  sp.  n. 
M .  montisarthuri  sp.  n. 
M.  tasmanicus  (Spencer,  1895). 

Other  species 

Sixty-five  Australian  species,  in  addition  to  the  Tasmanian  complement,  are 
listed  in  Jamieson,  igyic  (q.v.).  This  is  a  very  large  genus  with  many  species  in 
extra-Australian  parts  of  its  range. 

REMARKS.  Megascolex  is  a  polyphyletic  congeries  in  need  of  revision  and  con- 
stitutes one  of  the  major  problems  in  oligochaete  taxonomy  (Jamieson,  igyic).  It  is 
the  type-genus  of  the  tribe  Megascolecini,  purely  meronephric  megascolecines  in 
which,  if  there  is  a  median  stomate  nephridium,  this  opens  into  the  intestine. 

The  following  three  Tasmanian  species  are  provisionally  retained  in  Megascolex, 
with  the  definition  of  which  they  conform,  pending  revision  of  the  genus.  Sufficient 
grounds  exist  to  suggest  that  the  Tasmanian  species  comprise  one  or  more  separate 
genera  but  erection  of  the  latter  is  deferred  until  the  mainland  species  of  Megascolex 
are  revised. 

KEY  TO  TASMANIAN  SPECIES  OF  MEGASCOLEX 

1  Spermathecal  pores  2  pairs          ..........  2 

-  Spermathecal  pores  5  pairs.     Fig.  306  .  M.  tasmanicus  (Spencer,  1895) 

2  Spermathecal  pores  median  to  a,  almost  contiguous  ventrally.     Fig.  3oA 

M.  montisarthuri  sp.  n. 

-  Spermathecal  pores  in  ab.     Fig.  306    ......      M.  burniensis  sp.  n. 


Megascolex  burniensis  sp.  n. 

Fig.  4,  306,  32F 
1  =  65  mm,  w  =  4-0  mm,  s  =  92. 

Prostomium  tanylobous.  First  dorsal  pore  4/5.  Setae  12  on  XII ;  24  caudally 
(30  counted  in  some  segments)  ;  seta  become  irregular  in  the  posterior  25  segments  ; 
zz  =  zy  anteriorly  but  zz  =  2-4^  posteriorly  ;  aa  =  2-2'^ab  preclitellar,  aa  =  3-406 
postclitellar ;  aa  not  discernible  as  a  ventral  break  caudally.  Nephropores  not 
externally  visible.  Clitellum  annular  |XIII-XVII.  Male  pores  on  small  papillae 
in  ab,  nearer  a,  on  XVIII.  Genital  markings :  3  pairs  of  small  eye-like  markings 


320 


B.   G.   M.   JAMIESON 


sp.p 


FIG.  30.     Genital  fields.     A,  Megascolex  montisarthuri,  holotype.     B,  M.  burniensis, 
holotype.     C,  M.  tasmanicus,  specimen  i. 


TASMANIAN  EARTHWORMS  321 

with  centres  approximately  in  a,  one  pair  presetal  in  XVII,  the  other  two  pairs  in 
17/18  and  18/19  ;  two  similar  presetal  unpaired  markings  with  centres  in  a  in  XIX 
(right)  and  19/20  (left)  ;  two  larger  median  presetal  markings  present  in  XVIII  and 
XIX.  Female  pores  paired,  anterior  and  slightly  median  to  a  of  XIV.  Spermathe- 
cal  pores  2  pairs,  in  7/8  and  8/9,  on  small  papillae  slightly  lateral  of  a. 

Last  hearts  in  XIII  (hearts  in  X-XIII  latero-oesophageal  with  connective  to 
dorsal  and  supra-oesophageal  vessel)  ;  supra-oesophageal  in  VII-XIII,  well  deve- 
loped. Gizzard  large,  firm,  globular  with  anterior  rim,  in  V.  Extramural  calciferous 
glands  absent ;  oesophagus  with  circumferential  vascular  striae  in  VIII-XV  ; 
rugae  increasing  posteriorly,  especially  well  developed  in  XIV  and  XV.  Intestinal 
origin  XVII ;  typhlosole  absent.  Nephridia  poorly  preserved  ;  few  avesiculate, 
exonephric  meronephridia  on  each  side  in  each  segment ;  exonephric  tufts  in  II-VII ; 
caudally  exonephric  with  a  preseptal  funnel  on  each  side,  the  median  nephridium 
somewhat  enlarged.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ; 
seminal  vesicles  racemose  in  XI  and  XII.  Metagynous  ;  ovisacs  present  ?  (a  small, 
flocculent  mass,  on  the  right  side  only,  is  doubtfully  an  ovisac).  Prostates  flattened 
and  almost  square,  in  XVII  and  XVIII ;  truly  racemose,  the  muscular  duct  dividing 
into  three  immediately  within  the  gland ;  vas  deferens  joining  the  duct  near  its 
ental  end.  Penial  setae  absent.  Spermathecae  2  pairs,  in  VIII  and  IX,  diverti- 
culum  (inseminated)  single,  clavate,  uniloculate. 

MATERIAL  EXAMINED. 

Fern  Glade,  Emu  River,  Burnie,  I45°55'E.  4i°O5'S.  Map,  7/2,  Dr  J.  L.  Hickman,  24  Aug 
1954  -  holotype  TM  :  K/jSo. 

REMARKS.     The  genital  field  is  diagnostic  of  this  species. 


Megascolex  montisarthuri  sp.  n. 

Fig.  4,  soA,  326 
=  95.  105  mm,  s  =  3-6,  5-2  mm,  s  =  125,  123. 


Prostomium  canaliculate  or  not,  tanylobous  closed.  First  dorsal  pore  3/4,  4/5. 
Setae  18  (holotype),  12  (paratype  i)  in  XII,  caudally  with  24  or  more  in  a  segment ; 
rows  regular  in  the  fore-  and  midbody,  with  the  ventral  gap  =  $ab  and  much  wider 
dorsal  gap  ;  caudally  with  aa  unchanged  but  other  rows  irregular  and  no  appreciable 
dorsal  interruption  of  the  setal  circlet.  Nephropores  not  externally  visible.  Clitel- 
lum  annular,  XIII-XVII  (holotype,  paratype  i)  with  weak  development  in  XVIII 
(holotype).  Male  pores  almost  contiguous  midventrally  in  XVIII  on  a  pair  of 
united  papillae  which  lie  in  a  depression  with  tumid  borders  which  extends  almost 
from  the  setal  arc  of  XVII  to  that  of  XIX  and  laterally  reaches  b  lines  ;  a  midventral 
transverse  glandular  depression  with  tumid  margins  present  in  each  of  intersegments 
9/10,  16/17  an<i  I9/20»  spanning  cc  and  carrying  a  transverse  series  of  whitish  pore- 
like  stigmata.  Female  pores  anteromedian  to  setae  a  at  about  \aa.  Spermathecal 
pores  2  pairs,  on  medianly  closely  apposed  papillae  near  the  anterior  margins  of 
VIII  and  IX. 


322 


B.   G.  M.   JAMIESON 


TASMANIAN  EARTHWORMS  323 

Dorsal  vessel  single  ;  continuous  onto  the  pharynx  ;  last  hearts  in  XII  (hearts  in 
X-XII  latero-oesophageal,  with  connective  to  dorsal  and  supra-oesophageal  vessel)  ; 
supra-oesophageal  weakly  developed  ;  gizzard  very  well  developed,  in  V  but  extend- 
ing posteriorly  into  IX  by  displacement  of  attenuated  septa  ;  oesophagus  widened  to 
form  an  annular  calciferous  gland  with  numerous  high  radial  internal  lamellae  in  XV 
(holotype,  paratypes,  I,  8)  ;  similarly  developed  in  XVI  and  not  demarcated  from 
that  in  XV  in  holotype,  or  in  XIV  and  less  so  in  XIII  in  paratype  i ;  swollen  and 
vascular,  almost  reniform  on  each  side  in  XVII  in  paratypes  i  and  8.  Intestinal 
origin  XVIII  reaching  full  width  in  XIX  ;  typhlosole  strongly  developed ;  first 
appreciable  (rudimentary)  in  XX-XXI  (holotype,  paratypes  i,  8).  Nephridia 
several  exonephric,  astomate,  avesiculate  parietal  meronephridia  on  each  side  in 
each  segment ;  a  large  pair  of  tufts  in  III,  extending  into  II  and  IV  and  each  sending 
a  sheaf  of  ducts  (composite  duct)  to  enter  the  parietes  anterior  to  seta  b  of  II ; 
caudally  the  medianmost  nephridium  enlarged  as  a  megameronephridium  with  single 
preseptal  funnel  (exonephric)  (holotype).  Holandric  (funnels  iridescent)  ;  gymnor- 
chous  ;  seminal  vesicles  racemose,  in  XI  (small)  and  XII ;  metagynous  ;  small  ovi- 
sacs  in  XIV  ;  prostates  compact,  racemose,  restricted  to  XVIII  (holotype,  paratypes 
i,  8)  with  ramifying  internal  ducts  and  no  dominant  central  duct ;  vas  deferens 
joining  ectal  end  of  duct  (holotype).  Penial  setae  absent.  Spermathecae  2  pairs, 
in  VIII  and  IX  (the  ampullae  in  the  segment  anterior  to  its  pores  in  the  holotype 
only)  ;  diverticulum  (inseminated)  small,  clavate,  uniloculate  (holotype  ;  paratypes 
1,8). 

MATERIAL  EXAMINED. 

Mt  Arthur,  Weldborough  Pass,  1-6  miles  from  eastern  end,  i47°55'E.  4i°io'S.  Map,  9/3, 
Dr  J.  L.  Hickman,  26  Aug  1953  —  holotype  TM  :  K38i.  East  side  of  mountain  i47°2o'E. 
4i°i5'S.  Map,  9/1,  Mr  A.  J.  Dartnall  and  Mr  R.  C.  Kershaw,  15  Oct  1971  -paratypes  i,  8, 
16-18  BM(NH)  :  1973.2.48-52  ;  paratypes  2-7,  9-15  TM  :  1^382-394  ;  paratypes  19-25 
AM  :  W5332-5338  ;  paratypes  26-29  BJ  :  1:72-75. 

REMARKS.  The  closely  apposed  spermathecal  and  male  pores  and  the  very 
distinctive  accessory  genital  markings  permit  ready  identification  of  M.  montisarthuri 
and  jointly  distinguish  it  from  all  other  species. 

Of  29  paratypes  from  Mt  Arthur,  all  agree  with  the  holotype  in  possessing  the 
characteristic  median  transverse  genital  markings  at  16/17  and  !9/20,  the  median 
field  around  the  two  male  pores  being  variably  developed ;  25  of  these  have  an 
additional  transversely  punctate  genital  marking  extending  laterally  into  ab  in  9/10. 


FIG.  31.  Spermathecae  of  Cryptodrilus.  A,  C.  avesiculatus,  holotype  (right  IX).  B  &  C, 
C.  brunyensis  :  B,  holotype  (right  VIII)  ;  C,  paratype  (right  IX).  D,  C.  dartnalli, 
holotype  (left  IX).  E-G,  C.  enteronephricus  :  E,  holotype  (right  IX)  ;  F,  paratype  3 
(right  IX)  ;  G,  paratype  14  (right  IX^.  H,  C.  polynephricus  polynephricus,  specimen  12 
(right  IX).  I,  C.  polynephricus  ad  urethrae,  specimen  44  (right  IX).  J-M,  C.  p.  urethrae  : 
J,  holotype  (right  VIII)  ;  K,  paratype  i  (right  IX)  ;  L,  paratype  8  (right  IX)  ;  M, 
paratype  20  (right  IX) .  N  &  O,  C.  simsoni  :  N,  lectotype  (right  VI J  I)  ;  O,  specimen  2 
(right  IX).  All  except  L  to  the  scale  indicated. 


324  B.   G.  M.   JAMIESON 

Megascolex  tasmanicus  (Spencer,  1895) 
Fig.  4,  soC,  32H,  I 

Perichaeta  tasmanica  Spencer,  1895  :  47-48,  PI.  IV,  fig.  37-39. 
Megascolex  tasmanicus  ;  Michaelsen,  1900  :  217  ;   Jamieson,  19710  :  96. 

1  =  7°>  75  mm  (63-88  mm,  Spencer),  w  =  3-5  mm  (3-1  mm,  Spencer),  s  =  89,  93 
(specimens  i  and  2). 

Prostomium  epilobous  §  (^-|,  Spencer),  open  ;  peristomium  bifid  ventrally. 
First  dorsal  pore  4/5.  Setae  18  (specimen  i),  25  (specimen  2)  (20-22,  Spencer)  in 
XII  ;  28-30  caudally  (24-28,  Spencer)  ;  zz  =  2zy  anteriorly,  irregular  caudally, 
forming  a  definite  gap  in  the  setal  circlet  in  the  fore-  and  midbody  but  inappreciable 
at  the  caudal  extremity  ;  aa  =  2ab  anteriorly,  regular  throughout,  =  sab  caudally, 
a  definite  gap  throughout ;  setae  a  and  b  absent  in  XVIII.  Nephropores  not  ex- 
ternally visible.  Clitellum  weakly  developed,  annular,  in  XIII-XVII  (as  Spencer). 
Male  pores  small  but  conspicuous  transverse  slits  in  a  or  ab,  on  XVIII,  each  on  the 
suggestion  of  a  small  papilla,  in  a  circular,  glandular,  depressed  field.  Accessory 
genital  markings  incompletely  developed  but  well-defined,  presetal  median  trans- 
verse glandular  bands  in  IX,  X,  XI,  and  XII  extending  maximally  to  ab  (specimen  i) 
or  rudimentary  in  VIII-XI  (specimen  2)  ;  a  presetal,  median,  approximately  oval, 
depressed  glandular  patch  in  each  of  segments  XVII-XXII  ;  similar  but  paired 
circular  presetal  markings  present  in  each  of  segments  XX-XXIII,  almost  contiguous 
with  the  median  marking  where  one  occurs  in  the  same  segment  (markings  median  in 
IX,  X,  XI,  XX,  XXI  and  XXII,  Spencer).  Female  pores  a  pair  in  XIV,  shortly 
anterior  to  the  setal  arc  and  close  together  at  about  \aa  ;  spermathecal  pores  5  pairs 
of  small  but  distinct  white-rimmed  slits,  in  4/5-8/9,  in  a  lines. 

Internal  anatomy  (specimen  i)  :  dorsal  blood  vessel  single,  continuous  onto  the 
pharynx  ;  last  hearts  in  XII  (hearts  in  X-XII  latero-oesophageal,  each  with  con- 
nective to  dorsal  and  supra-oesophageal  vessel)  ;  supra-oesophageal  in  IX-|XIII, 
well  developed.  Gizzard  muscular,  but  small  and  easily  compressed,  in  V  ;  true 
ovoidal  calciferous  glands,  3  pairs  in  segments  X,  XI  and  XII,  each  sessile,  with  a 
narrow  connection  with  the  lateral  wall  of  the  oesophagus,  the  lumen  of  the  gland 
almost  occluded  by  approximately  12  laminae  which  converge  on  the  connection 
with  the  oesophagus  but  do  not  unite  ;  intestinal  origin  XV  (XVIII,  Spencer)  ;  a 
very  low  but  definite  typhlosole  present,  first  appreciable,  though  very  rudimentary, 


FIG.  32.  Spermathecae.  A,  Graliophilus  tripapillatus,  holotype  (right  IX).  B,  Pinguidri- 
lus  tasmanicus,  lectotype  (right  IX).  C  &  D,  Hickmaniella  opisthogaster,  holotype  :  C, 
ventral  view  ;  D,  dorsal  view  (right  IX).  E,  Rhododrilus  littoralis,  holotype  (right  IX). 
F,  Megascolex  burniensis,  holotype  (right  IX).  G,  M.  montisarthuri,  holotype  (right  IX). 
H  &  I,  M.  tasmanicus,  specimen  i  :  H,  left  IX  ;  I,  right  VIII.  J,  Oreoscolex  bidiverti- 
culatus,  holotype  (right  IX).  K,  O.  campestris,  specimen  i  (right  IX).  L,  O.  irregularis, 
specimen  I  (right  VIII).  M,  O.  peculiaris,  holotype  (right  IX).  N  &  O,  O.  longus,  holo- 
type :  N,  dorsal  view  ;  O,  ventral  view  (right  IX).  P  &  Q,  O.  sexthecatus,  holotype  : 
P,  ventral  view  ;  Q,  dorsal  view  (right  VIII).  R,  0.  wellingtonensis,  specimen  i  (right  IX). 
S,  Pseudocryptodrilus  acanthodriloides,  holotype  (right  IX).  All  to  the  same  scale 
excepting  E  which  is  twice  the  scale  indicated. 


TASMANIAN  EARTHWORMS 


325 


326  B.   G.  M.   JAMIESON 

in  XVI ;  caeca  and  muscular  thickening  absent.  A  few  small  exonephric  tufted 
nephridia  on  each  side  in  IV,  small  nephridia  apparently  present  in  III  (and  II  ?)  ; 
astomate,  avesiculate,  exonephric  micromeronephridia  numerous  on  each  side,  form- 
ing a  dense  parietal  band,  in  XV  posteriorly  ;  less  numerous  in  V-XIV.  Caudally 
with  numerous  micromeronephridia  and  a  median  nephridium  which  has  several 
spiral  loops  and  might  be  considered  a  megameronephridium.  The  median  nephri- 
dium with  a  large  preseptal  funnel  and  with  its  duct  joining  a  paired  duct  running 
longitudinally  on  the  roof  of  the  intestine  shortly  lateral  of  the  dorsal  blood  vessel ; 
this  longitudinal  duct  connected  to  and  presumably  opening  into  the  intestine 
posteriorly  in  each  segment.  Caudal  micromeronephridia  associated  with  preseptal 
bodies  which  may  be  reduced  funnels  ;  whether  enteronephric  or  exonephric  not 
determined.  Holandric  (funnels  iridescent  in  X  and  XI)  ;  gymnorchous  ;  seminal 
vesicles  racemose,  very  large,  in  IX  and  XII,  the  larger  in  IX.  Metagynous  ; 
ovisacs  present.  Prostates  racemose,  bipartite,  one  lobe  in  XVIII,  the  other  in  XIX, 
the  muscular  duct  bifurcating  immediately  within  the  gland,  joined  entally  by  the 
vas  deferens.  Penial  setae  absent.  Spermathecae  5  pairs,  diverticulum  single, 
longer  than  the  ampulla,  clavate,  uniloculate. 

TYPE-LOCALITY.    Emu  Bay,  Tasmania,  and  King  Island  in  Bass  Strait.    Map,  7/2. 
MATERIAL  EXAMINED. 

Table  Cape,  I45°45'E.  4i°oo'S.  Map,  7/1,  Dr  J.  L.  Hickman,  24  Aug  1954  -  specimen  * 
TM  :  K395  ;  specimen  2  BJ  :  Tj6. 

REMARKS.  M.  tasmanicus  is  questionably  distinct  from  the  Victorian  species 
M.  hoggi  (Spencer  i8g2b)  and  its  occurrence  on  King  Island  may  indicate  that 
it  represents  a  Tasmanian  population  of  the  latter  species.  If  so,  it  would  con- 
stitute the  only  known  case  of  a  megascolecid  species  shared  between  Tasmania 
and  the  mainland.  Spencer  (1895)  drew  attention  to  the  similarity  of  M. 
tasmanicus,  M.  sylvaticus,  M.  hoggi,  M.  halli,  M.  rubra,  M.  frenchi  and  M.  steeli,  all 
Victorian  species  of  Spencer  (i8g2b),  in  having  a  bifid  peristomium,  5  pairs  of  sper- 
mathecal  pores,  calciferous  glands  in  X-XII  and  bilobed  prostates.  This  combina- 
tion of  characters  suggests  a  basis  for  recognition  of  a  new  (wholly  dichogastrin  ?) 
genus  distinct  from  Megascolex. 

ACKNOWLEDGEMENTS 

The  author  is  deeply  indebted  to  Dr  J.  L.  Hickman  of  the  University  of  Tasmania 
for  providing  his  incomparable  collections  and  to  Mr  R.  W.  Sims  for  providing 
facilities  in  the  British  Museum  (Natural  History)  for  this  study.  Thanks  are  also  due 
to  other  collectors,  acknowledged  in  the  text,  and  to  Miss  E.  Pope,  Dr  P.  Hutchings, 
Mr  A.  J.  Dartnall  and  Dr  B.  J.  Smith  of  the  Australian,  Tasmanian  and  National 
Museums  for  loan  of  material.  Mr  E.  A.  Bradbury,  Mr  T.  Walker  and  Mr  J.  Casey  are 
thanked  for  their  assistance  and  Miss  O.  Page-Hanify  for  her  efficient  typing.  Mr 
Walker  took  the  photographs  in  cooperation  with  the  Electron  Microscope  Centre, 
University  of  Queensland,  and  also  prepared  the  maps.  All  line  drawings  are  by  the 
author.  The  work  was  made  possible  by  Australian  Research  Grants  Committee 
grant  nos.  23926o-300-R-Zool-ARGC-i2O-7i-72. 


TASMANIAN  EARTHWORMS  327 

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zool.  Soc.  Lond.  1932  :  899-941. 

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WALLACE,  C.  C.     1972.     An  examination  of  the  classification  of  some  Australian  megascolecid 

earthworms  (Annelida  :  Oligochaeta)  by  numerical  methods.      Mem.  Qd  Mus.  16  (2)  :  191- 

209. 

BARRIE  G.  M.  JAMIESON,  Ph.D. 

Department  of  Zoology 

UNIVERSITY  OF  QUEENSLAND 

ST  LUCIA 

BRISBANE  4067 

QUEENSLAND 

AUSTRALIA 


PLATES    1-12 


Scanning  electron  micrographs  of  penial  setae,     i,  Graliophilus  tripapillatus,  holotype. 

2  &  3,  Cryptodrilus  avesiculatus,  holotype. 

4-7,  C.  brunyensis  :   4-6,  one  seta  ;    7,  a  second  seta  of  paratype  i. 

8-12,  C.  dartnalli  :   8-10,  paratype  8  ;    u  &  12,  holotype. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES    1-12 


PLATES    13-24 

Scanning  electron  micrographs  of  penial  setae.  13-19,  Cryptodrilus  enter onephricus  :  13-15, 
paratype  i  ;  16-19,  paratype  14. 

20-24,  C.  polynephricus  polynephricus  :  20  &  21,  specimen  3  ;  22,  specimen  i  ;  23  &  24, 
specimen  5. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES    13-24 


PLATES   25-36 

Scanning   electron   micrographs   of   penial   setae.     25-31,    Cryptodrilus  polynephricus  poly- 
nephricus  :    25,  specimen  10  ;    26,  specimen  8  ;    27  &  28,  specimen  24  ;    29-31,  specimen  34. 
32-36,  C.  p.  polynephricus  ad  urethrae  :   32  &  33,  specimen  42  ;    34-36,  specimen  44. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   25-36 


36 


PLATES   37-48 

Scanning  electron   micrographs  of  penial   setae.     37-43,    Cryptodrihis  polynephricus  poly- 
nephricus  ad  urethrae  :   37-39,  specimen  44  ;   40-42,  specimen  48  ;    43,  specimen  49. 
44-48,  C.  p.  urethrae  :   44-47,  holotype  ;   48,  paratype  6. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   37-48 


PLATES   49-60 

Scanning  electron  micrographs  of  penial  setae.  49-59,  Cryptodrilus  polynephricus  urethrae  : 
49  &  50,  second  seta,  paratype  6  ;  51  &  52,  paratype  20  ;  53-59,  3  penial  setae  of  paratype  i, 
53,  lateral  view  of  a  seta  ;  54,  ventral  view  of  same  ;  55,  dorsal  view  of  same  ;  56  &  57,  lateral 
and  dorsal  detail  ;  58,  lateral  view  of  second  seta  ;  59,  detail  of  third  seta. 

60,  Rhododrilus  littoralis,  paratype  i. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   49-60 


SO 


PLATES   61-72 

Scanning  electron  micrographs  of  penial  setae.     61-63,  Pseudocryptodrihis  acanthodriloides, 
paratype  3. 

64-66,  Hickmaniella  opisthogaster  :   64,  holotype  ;    65  &  66,  paratype  i. 

67,  Oreoscolex  bidiverticulatus,  holotype. 

68-72,  O.  campestris  :   68  &  69,  specimen  i  ;    70-72,  specimen  2. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   61-72 


PLATES   73-84 

Scanning  electron  micrographs  of  penial  setae.     73-80,  Oreoscolex  longus  :    73-75,  holotype 
76-78,  paratype  i  ;    79  &  80,  paratype  7. 
81-82,  O.  peculiaris,  holotype. 
83  &  84,  O.  sexthecatus,  holotype. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES 


PLATES   85-90 

Scanning  electron  micrographs  of  penial  setae.     85  &  86,  Oreoscolex  sexthecatus,  seta  shown  in 
plate  83,  holotype. 

87-90,  O.  wellingtonensis ,  specimen  i. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   85-90 


90 


PLATES   91-96 

Scanning  electron  micrographs  of  penial  setae.     91  &  92,  Perionychella  (P.)  capensis,  paratype  i . 
93,  P.  (Vesiculodrilus]  bithecata,  paratype  3.     94-96,  P.  (V .)  dilwynnia  :    94  &  95,  Butler's 
Gorge  Road  specimen  ;    96,  Tarraleah  pipeline  specimen. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   91-96 


PLATES   97-102 

Scanning  electron  micrographs  of  penial  setae.  97  &  98,  Perionychella  (Vesiculodrilus) 
hobartensis  :  97,  14/6,  BM(NH)  :  1972.8.15  ;  98,  14/5,  TM  :  1^304. 

99-102,  P.  (V .)  mortoni  :  99  &  100,  left  and  right  setae  respectively  of  East  Risclon  specimen  ; 
1 01  &  1 02,  specimen  described  from  Shoobridge  Bend. 


Bull.  Br.  Mus.  nat.  Hist.  (Zool.)  26,  4 


PLATES   97-102 


1O1 


1O2 


A  LIST  OF  SUPPLEMENTS 
TO  THE  ZOOLOGICAL  SERIES 

OF  THE  BULLETIN  OF 
THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 


1.  KAY,  E.  ALISON.     Marine  Molluscs  in  the  Cuming  Collection  British  Museum 
(Natural  History)  described  by  William  Harper  Pease.     Pp.  96  ;    14  Plates. 
1965.     (Out  of  Print.)     £3.75. 

2.  WHITEHEAD,  P.  J.  P.    The  Clupeoid  Fishes  described  by  Lacepede,  Cuvier  and 
Valenciennes.     Pp.  180  ;   n  Plates,  15  Text-figures.     1967.     £4. 

3.  TAYLOR,  J.  D.,  KENNEDY,  W.  J.  &  HALL,  A.  The  Shell  Structure  and  Mineralogy 
of  the  Bivalvia.      Introduction.      Nuculacea-Trigonacea.      Pp.  125  ;  29  Plates 
77  Text-figures.     1969.    £4.50. 

4.  HAYNES,  J.  R.     Cardigan  Bay  Recent  Foraminifera  (Cruises  of  the  R.V.  Antur) 
1962-1964.     Pp.  245  ;  33  Plates,  47  Text-figures.     1973.     £10.80. 

5.  WHITEHEAD,  P.  J.  P.     The  Clupeoid  Fishes  of  the  Guianas.     Pp.  227  ;    72 
Text-figures.     1973.     £9.70. 


Printed  in  Great  Britain  by  John  Wright  and  Sons  Ltd.  at  The  Stonebridge  Press,  Bristol  BS4  jNU 


22  JUL1974 


THE  FRESHWATER  FISHES  OF 

MUNGO  AND  MEME  AND  LAKES 

KOTTO,  MBOANDONG  AND 

SODEN,  WEST  CAMEROON 


E.  TREWAVAS 


BULLETIN  OF 

THE    BRITISH    MUSEUM    (NATURAL  HISTORY) 
ZOOLOGY  Vol.  26  No.  5 

LONDON:  1974 


22  JULI9! 


THE  FRESHWATER  FISHES   OF  RIVERS 

MUNGO  AND  MEME  AND  LAKES 

KOTTO,  MBOANDONG  AND 

SODEN,  WEST  CAMEROON 


BY 

ETHELWYNN  TREWAVAS 


Pp  329-419;  5  Plates,  17  Text-figures 


BULLETIN  OF 

THE  BRITISH  MUSEUM  (NATURAL  HISTORY) 
ZOOLOGY  Vol.  26  No.  5 

LONDON:  1974 


THE      BULLETIN      OF     THE     BRITISH     MUSEUM 

(NATURAL  HISTORY),  instituted  in  1949,  is 
issued  in  five  series  corresponding  to  the  Departments 
of  the  Museum,  and  an  Historical  series. 

Parts  will  appear  at  irregular  intervals  as  they 
become  ready.  Volumes  will  contain  about  three  or 
four  hundred  pages,  and  will  not  necessarily  be 
completed  within  one  calendar  year. 

In  1965  a  separate  supplementary  series  of  longer 
papers  was  instituted,  numbered  serially  for  each 
Department. 

This  paper  is  Vol.  26,  No.  5,  of  the  Zoological  series. 
The  abbreviated  titles  of  periodicals  cited  follow  those 
of  the  World  List  of  Scientific  Periodicals. 


World  List  abbreviation  : 
Bull.  Br.  Mus.  nat.  Hist.  (Zool.) 


Trustees  of  the  British  Museum  (Natural  History),  1974 


TRUSTEES   OF 
THE  BRITISH  MUSEUM   (NATURAL  HISTORY) 

Issued  24  May,  1974  Price  £5.60 


THE  FRESHWATER  FISHES  OF  RIVERS 

MUNGO  AND  MEME  AND  LAKES 

KOTTO,  MBOANDONG  AND 

SODEN,  WEST  CAMEROON 

By  ETHELWYNN  TREWAVAS 

CONTENTS 


SYNOPSIS  . 
INTRODUCTION 


Page 
332 
332 


DESCRIPTION  OF  THE  LOCALITIES  .......  333 

ABBREVIATIONS  AND  MEASUREMENTS  ......  333 

MORMYRIDAE  :   Mormyrus  tapirus         .......  334 

Brienomyrus  brachyistius       ......  335 

CHARACIDAE  :  Alestes  macrolepidotus   .......  339 

Alestes  longipinnis         .......  339 

CYPRINIDAE  :  Labeo  camerunensis  n.  sp.        ......  340 

Barbus  batesii        ........  343 

Barbus  mungoensis  n.  sp.         ......  345 

Barbus  callipterus.          .......  347 

Barbus  camptacanthus    .......  349 

Barbus  thysi  n.  sp.          .......  350 

CLARIIDAE  :   Clarias  walkeri         ........  356 

BAGRIDAE  :   Auchenoglanis  ahli  .......  356 

MOCHOKIDAE  :   Chiloglanis  :    structure  of  head      .....  358 

Chiloglanis  tnicropogon  .          .          .          .          .          .  361 

Chiloglanis  disneyi  n.  sp.        ......  365 

MALAPTERURIDAE  :   Malapterurus  electricus  ......  367 

CYPRINODONTIDAE  :  Epiplatys  sexfasciatus  .          .          .          .          .          .  367 

Aphyosemion  bivittatum         .....  370 

Aphyosemion  celiae        .          .          .          .          .          .  371 

Aphyosemion  oeseri       .          .          .          .          .          .  373 

Procatopus :  specific  characters  and  sexual  dimorphism  376 

Procatopus  similis         ......  384 

Procatopus  nototaenia    ......  385 

Procatopus  glaucicaudis          .          .          .          .          .  386 

Procatopus  lacustris  n.  sp.      .....  386 

CICHLIDAE  :   Hemichromis  elongatus      .......  387 

Chromidotilapia  :   definition      ......  388 

Feeding  mechanism  :   contrast  with  Tilapia    .          .          .  389 

Chromidotilapia  finleyi  n.  sp.  .....  393 

Chromidotilapia  guntheri  guntheri  and  C.  guntheri  loennbergi  398 

Tilapia  camerunensis        .......  402 

Tilapia  kotta'e  ........  405 

Tilapia  mariae         ........  405 

Sarotherodon  galilaeus       .......  408 

ZOOGEOGRAPHICAL  SUMMARY  AND  CONCLUSIONS     .....  410 

ACKNOWLEDGEMENTS          .........  414 

REFERENCES     ...........  414 


332  E.   TREWAVAS 

SYNOPSIS 

Collections  of  fishes  from  the  upper  Mungo  and  its  tributaries  and  from  Lakes  Kotto,  Mboan- 
dong  and  Soden  are  described  and  their  relationships  discussed.  The  status  of  Brienomyrus 
longianalis  is  examined  in  the  light  of  variation  within  and  between  populations  of  B.  brachyis- 
tius.  A  new  species  of  Labeo  is  described.  The  Mungo  contains  two  species  of  Barbus  with 
parallel-striate  scales,  one  of  which  is  given  a  new  name.  The  three  small  Barbus  species  with 
radiating  striae  on  the  scales  are  identical  with  those  of  Fernando  Poo  and  one  of  them,  here 
given  a  new  name,  is  so  far  reported  only  from  Fernando  Poo  and  the  Mungo  and  Meme  systems. 
In  the  course  of  comparisons  with  this  species  records  of  the  geographical  distribution  of  B. 
trispilos  and  B.  liberiensis  are  critically  examined  and  possible  synonyms  of  B.  baudoni  are 
discussed. 

Specimens  of  Auchenoglanis  from  two  localities  prompt  questions,  but  no  answers,  concerning 
the  specific  characters  within  this  genus.  Two  species  of  Chiloglanis  were  collected  in  the  stony 
beds  of  swift  streams.  One  of  them  is  described  as  new  and  some  special  features  in  the  structure 
of  the  head  in  this  genus  are  described. 

Seven  species  of  cyprinodonts  are  recognized  in  the  area,  one,  a  Procatopus  of  Lake  Soden,  new. 
The  identity  and  synonymy  of  Aphyosemion  oeseri  Schmidt  are  discussed  and  a  new  description 
is  given  of  Procatopus  similis  Ahl. 

The  structure  of  the  pharynx  in  Chromidotilapia  is  described,  C.  loennbergi  is  considered  to  be  a 
subspecies  of  C.  guntheri,  a  West  African  species  present  also  in  the  Mungo  together  with  a  new 
species  related  to  C.  batesii.  The  new  species  is  described  and  compared  with  samples  from 
Fernando  Poo  and  South  Cameroun.  The  Hemichromis  of  the  area  is  the  'B  form  of  H.  fasciatus', 
here  tentatively  identified  with  H.  elongatus  Guichenot. 

The  description  of  Tilapia  kottae  is  amplified  and  the  name  T.  camerunensis  Lonnberg  is  revived 
for  a  related  species  inhabiting  the  Mungo  as  well  as  the  Meme  and  distinct  from  T.  cameronensis 
Holly  of  the  Sanaga  system.  T.  mariae  of  Lake  Kotto  is  no  longer  considered  to  merit  sub- 
specific  rank.  Some  notes  on  Sarotherodon  galilaeus  and  its  subspecies  are  included. 

The  Mungo-Meme  fish  fauna  is  seen  as  part  of  that  of  the  forest  zone  from  western  Nigeria  to 
South  Cameroun,  having  none  but  the  most  widespread  species  in  common  with  the  Chad  basin. 
Lakes  Kotto  and  Mboandong  contain  a  slightly  modified  section  of  this  fauna  and  Fernando  Poo 
harbours  twelve  species  common  to  it  and  the  Mungo  and  Meme  systems,  adapted  to  life  in  clear 
running  water.  This  throws  into  greater  relief  the  high  degree  of  endemism  in  the  crater  lake 
Barombi  Mbo. 


INTRODUCTION 

A  PREVIOUS  paper  (Trewavas,  Green  &  Corbet,  1972)  dealt  with  the  taxonomy  and 
ecology  of  the  fishes  of  Barombi  Mbo,  results  of  an  expedition  by  three  members  of 
Westfield  College,  University  of  London,  and  myself  in  March-April  1970.  The 
present  paper  describes  the  fishes  collected  during  that  expedition  in  the  other  waters 
visited.  The  ecology  of  those  of  Kotto  and  Mboandong  has  already  formed  the 
subject  matter  of  a  report  by  Corbet  et  al.  (1973). 

We  were  interested  to  get  good  samples  from  R.  Mungo  and  its  tributaries  and 
from  the  outflow  of  Barombi  Kotto  which  belongs  to  the  Meme  system,  as  a  back- 
ground to  the  endemism  in  the  lakes,  especially  Barombi  Mbo  ;  and  we  wished  to  get 
further  light  on  the  relationships  of  the  Kotto  fishes. 

For  the  Mungo  collections  we  are  entirely  indebted  to  Dr  Henry  Disney,  who  set 
narrow-meshed  traps  overnight  and  used  a  hand-net  to  sample  the  river  and  its 
tributaries  at  and  above  the  bridge  near  Etam  in  the  course  of  his  field  studies  of 
Simulium  as  a  vector  of  Onchocerca.  When  we  did  not  accompany  him  he  brought 


FISHES   OF  WEST  CAMEROON  333 

living  fishes  to  our  headquarters  near  the  lake  and  he  made  further  collections  in 
October  and  November,  1970. 

Kotto,  Mboandong  and  their  neighbouring  streams  were  sampled  by  our  colleague 
Dr  John  Griffith  by  means  of  basket  traps  and  fyke  net  and  by  Professor  Green  and 
Dr  Corbet  with  hand-nets.  In  addition  we  purchased  fish  from  cast-net  fishermen 
and  boys  with  hand-lines. 

Comparisons  of  some  species  with  their  representatives  in  other  waters  are  under- 
taken as  a  contribution  to  West  African  ichthyogeography. 


DESCRIPTION  OF  THE  LOCALITIES 

The  lakes  have  been  described  (Trewavas,  1962  ;  Corbet  et  al.  1973).  The 
whole  area  is  in  the  forest  zone  of  the  volcanic  hills  around  Mount  Cameroon. 

R.  Mungo  at  the  bridge  near  Etam  is  about  a  quarter  of  a  mile  broad.  There  is  a 
waterfall  not  far  above  the  bridge  and  the  current  is  rapid.  At  our  early  visits 
people  could  wade  to  the  sandbanks  exposed  in  its  bed,  but  before  we  left  heavy  rain 
had  swollen  it  so  that  the  sandbanks  were  submerged  in  a  broad  brown  flood. 

Between  Baduma  and  Bolo  the  river  was  much  narrower,  clear  and  shallow,  and 
the  same  description  applies  to  tributaries  from  the  Blackwater  upwards.  The  bed 
of  the  Mangusu  was  dry  at  the  beginning  of  our  visit,  but  at  the  end  the  stream  was  in 
spate  and  yielded  a  cyprinodont  species. 

In  the  Kotto  area,  R.  Nganjoke,  the  outlet  of  Lake  Kotto  flowing  to  the  Meme, 
was  sampled  where  it  had  become  a  clear  shallow  stream  with  rocky  bed,  in  our  time 
crossed  by  stepping  stones  but  since  bridged.  R.  Nyoke  (or  Njoke),  about  a  mile 
S.E.  of  Lake  Kotto,  is  not  connected  with  the  lake  but  is  a  tributary  of  the  lower 
Mungo  (information  from  Dr  Disney).  The  water  is  clear  and  is  used  by  the  villagers 
for  drinking  if,  as  rarely  happens,  there  has  not  been  enough  rain  to  be  collected  for 
that  purpose. 

The  Barombis  living  on  the  island  in  Lake  Kotto  obtain  their  drinking  water  from 
clear  streams  (called  Tungs)  that  enter  the  lake  from  the  crater  rim.  In  these  we 
caught  cyprinodonts  and  Barbus  callipterus. 


ABBREVIATIONS  AND  MEASUREMENTS 

BMNH  =  British  Museum  (Natural  History). 

SL  =  Standard  Length,  i.e.  excluding  the  caudal  fin  and  (except  in  Procatopus) 
excluding  any  projecting  part  of  the  lower  jaw. 

Interorbital  width  in  Ostariophysi  includes  the  skinny  rim  of  the  orbit  (which 
covers  a  flexible  bony  flange).  In  Cichlidae  the  callipers  are  pressed  against  the 
skin  so  that  this  measurement  is  virtually  that  of  the  bony  interorbital  part  of  the 
roof  of  the  skull. 

Depth  of  preorbital  in  Cichlidae  is  the  depth  of  the  preorbital  or  lacrimal  bone 
measured  from  the  middle  of  its  orbital  edge  along  a  line  continuing  the  radius  of 
the  eye. 


334 


E.   TREWAVAS 


FIG.  i.  Sketch  map  of  the  waters  of  West  Cameroon  in  which  the  collections  were 
made.  Drawn  by  J.  Green  from  Disney,  1971.  The  stream  Nyoke  is  the  unlabelled  one 
crossing  the  'K'  of  Kotto. 


MORMYRIDAE 
Mormyrus  tapirus  Pappenheim,  1905 

Pappenheim,  1905  :  217  ;  id.  1907  :  362  pi.  xiii  fig.  3  (R.  Lokundje  =  Lokoundie,  southern 
Cameroun)  ;  Boulenger,  1909:  133  fig.  in  ;  Holly,  1927:  120  (Nachtigal  Rapids  and  R. 
Mbam,  Sanaga  system)  ;  id.  1930  :  225. 

Mormyrus  goheeni  (Fowler  ?)  ;  Pellegrin,  1927  :  295  (R.  Nyong)  ;  Holly,  1930  :  225  (from 
Pellegrin) . 

Three  specimens  collected  by  Dr  Disney  in  R.  Wo  we,  October,  1970,  respectively 
132,  170  and  173  mm  in  SL. 


FISHES   OF  WEST   CAMEROON  335 

These  are  a  little  smaller  than  most  previously  described  specimens,  so  a  descrip- 
tion is  not  superfluous. 

Depth  of  body  20-23-5%  SL,  length  of  head  27-6-28-6,  length  of  pectoral  fin 
17-1-17-4,  of  pelvic  10-8-12-7,  of  dorsal  base  48-0-48-5,  of  anal  base  16-9-18-3. 
Length  of  caudal  peduncle  12-9-13-6%  SL,  2-8-3-0  times  its  depth. 

Snout  narrow  and  curved  downwards  to  end  in  the  thick-lipped  mouth,  its  length 
41-48-5%  length  of  head,  0-9-1-12  times  postocular  part  of  head.  Diameter  of  eye 
14-5-20-6%  length  of  head. 

Scales  84-98  in  the  lateral  line,  about  14-17  between  lateral  line  and  origin  of 
dorsal,  8-12  from  lateral  line  to  origin  of  anal,  13-16  around  caudal  peduncle. 

Dorsal  62-66,  the  first  ray  minute,  the  next  two  simple.  Anal  24-26,  2  or  3  of 
which  are  simple. 

Teeth  3  in  upper  jaw,  6  in  lower  (but  2  +  3  lower  in  one,  the  third  of  the  right  side 
being  lost  or  not  yet  formed). 

The  allometries  revealed  in  comparison  with  earlier  descriptions  are  the  usual 
negative  allometry  of  the  eye  and  the  strong  positive  allometry  of  the  snout.  Holly 
has  measured  this  in  terms  of  the  eye-diameter,  thus  emphasizing  the  change,  but 
showing  continuation  of  the  trend  observable  even  within  the  narrow  size-range  of  our 
sample. 

Our  specimens  have  fewer  teeth  in  the  upper  jaw  than  those  formerly  described 
(Pappenheim,  Boulenger  and  Holly  5,  Fowler  6  in  M.  goheeni,  the  last  perhaps  with- 
out realizing  the  usual  presence  of  a  median  tooth  in  Mormyridae).  This  may  also 
be  related  to  size.  Pappenheim  and  Boulenger  found  an  additional  tooth  on  each 
side  too  in  the  lower  jaw,  making  a  total  of  8. 

Holly  has  already  found  in  7  specimens  from  Rivers  Lokundje  and  Sanaga  a 
range  of  60-73  rays  in  the  dorsal  fin.  It  may  have  been  from  specimens  at  the  lower 
end  of  this  range  that  Pellegrin  identified  his  Nyong  specimens  as  M.  goheeni  (Fowler 
gave  60,  58  and  61  for  the  type  and  two  other  Liberian  specimens).  But  to  my  mind 
the  presence  of  low  numbers  in  samples  from  Cameroun  should  rather  be  taken  as 
evidence  that  there  is  one  species  from  Liberia  to  Cameroun  than  that  two  species 
so  defined  are  present  in  Cameroun. 

The  type  of  M.  goheeni  Fowler  (1919  ;  Liberia)  was  described  as  having  80  scales 
in  the  lateral  line.  The  anterior  scales  are  difficult  to  count  because  of  the  thickened 
skin  overlying  them  and  some  irregularity  in  their  alignment.  The  Mungo  counts 
not  only  include  Pappenheim's  and  Boulenger's  count  of  90,  but  bring  the  range 
nearer  to  Fowler's  datum  and  make  it  more  probable  that  M .  goheeni  is  a  synonym 
of  M .  tapirus.  There  are  no  records  under  either  name,  to  my  knowledge,  between 
Liberia  and  Cameroun,  and  since  Fowler's  description  of  the  types  of  M.  goheeni  no 
further  records  from  Liberia. 

BRIENOMYRUS  Taverne,  1971 
Taverne,  igyia  :  101  and  108  ;   id.  igyib.     Type  species  Marcusenius  brachyistius  Gill. 

Taverne  shows  that  this  genus  is  distinguished  from  Marcusenius  Gill  (type  species 
Mormyrus  cyprinoides  Linnaeus)  by  the  broader,  curved  mesethmoid,  the  absence  of 


336  E.   TREWAVAS 

lateral  ethmoids  and  the  presence  of  6  circumorbital  bones,  the  antorbital  being  free 
from  the  first  infraorbital. 


Brienomyrus  brachyistius  (Gill) 

Marcusenius  brachyistius  Gill,  1862  :  139  (Liberia  ?)  ;  Boulenger,  1909  :  67  fig.  50  ;  id.  1916  :  158  ; 

Pellegrin,  1927  :  295  (Wuri  and  Nyong)  ;   Holly,  1930  :  222. 
Mormyrus  microcephalus  Giinther,  1867  '.115  (Ogowe). 
Marcusenius  liberiensis  Steindachner,  1894  :  67  (Liberia). 
?  Marcusenius  longianalis  Boulenger,  igoia  :  5  pi.  iii  fig.  i  (Niger  delta)  id.  1909  :  69  fig.  51  ; 

id.  1916  :  159. 

?  Marcusenius  adustus  Fowler,  1936  :  247  fig.  4  (nr  Kribi,  Cameroun). 
?  Marcusenius  sp.  Reed,  1967  :  26  fig.  33. 

Our  specimens  are  : 

SL  (mm)  Locality  Collector 

4  121-176  R.  Mungo 

between  Baduma  and  Bolo  R.  H.  L.  Disney 

2      79*5-133-5        R.  Mungo  nr  Etam  R.  H.  L.  Disney 

5  95'5-*37'5        R-  Wowe  R.  H.  L.  Disney 
i     163                      R.  Nganjoke  J.  Griffith 

The  holotype,  of  unknown  locality,  'probably  Liberia',  had  17  dorsal  and  27  anal 
rays.  In  Cameroun  the  species  has  been  recorded  from  R.  Nyong  at  Dehane,  R. 
Wuri  at  Yabassi  (Pellegrin,  1927),  Kribi  (Boulenger,  1916  :  158),  Tiko  near  the  coast 
in  the  R.  Matute  N.  of  the  Mungo  mouth  (Trewavas,  1962). 

Our  Nganjoke  fish  is  slightly  deeper-bodied  than  all  but  the  smallest  of  the  Mungo 
specimens  and  has  a  deeper  caudal  peduncle.  It  resembles  Boulenger's  figure  of  B. 
brachyistius. 

The  ii  Mungo  specimens  are  all  slender  fishes,  only  the  smallest  having  a  maximum 
depth  of  more  than  18%  SL  and  only  one  having  a  caudal  peduncle  less  than  3  times 
as  long  as  deep.  This  led  me  to  compare  them  not  only  with  B.  brachyistius  but  also 
with  B.  longianalis  of  the  lower  Niger. 

The  results,  summarized  in  Tables  i  and  2,  make  it  difficult  to  maintain  B. 
longianalis  as  a  separate  species.  According  to  Boulenger's  key  (1909  :  61)  the  only 
trenchant  contrast  is  in  the  number  of  anal  rays  (31-33  v.  25-30).  But  one  of  the 
syntypes  of  M.  microcephalus  Giinther  (R.  Ogowe),  synonymized  by  Boulenger  with 
brachyistius,  also  has  31  anal  rays  and  6  other  specimens  from  the  Lower  Niger, 
included  in  longianalis,  have  30-32.  It  is  no  longer  justifiable  therefore  to  draw 
an  arbitrary  line  between  30  and  31  rays.  It  is  nevertheless  true  that  in  the  lower 
Niger  long  anals  are  associated  with  low  numbers  of  dorsal  rays  (Table  3).  Else- 
where high  anal  counts  are  associated  with  rather  high  numbers  of  dorsal  rays  (Sierra 
Leone  and  R.  Gambia)  or  low  dorsal  counts  with  low  numbers  of  anal  rays  (Niari- 
Kouilou,  Chiloango  and  Congo)  (Table  3).  In  the  Mungo  the  slender  shape  is  as- 
sociated with  intermediate  numbers  of  dorsal  and  anal  rays,  although  the  anal 
numbers  range  higher  than  in  the  presumed  type  locality  (Liberia). 


FISHES   OF  WEST  CAMEROON  337 

TABLE  i 

Some  critical  characters  in  samples  of  Brienomyrus  brachyistius  and  possible  synonyms 


n 

D 

A 

A/D 

sc. 

depth 

c.p.l/d 

vert. 

Liberia 

10 

17-18 

24-27 

1-5-2-0 

56-63 

20-6-24-8 

2-2-2-9 

44-45 

Ghana  and  nr 

Lagos 

5 

15-16 

27-28 

1-95-2-25 

- 

17-2-22-5 

2-6-3-3 

44-45 

Nr  Oban, 

Calabar 

3 

16-17 

28-29 

1-9-2-2 

- 

18-5-20-6 

2-6-3-1 

45.  46 

Mungo 

ii 

16-18 

27-29 

1-6-2-0 

59-65 

14-2-20-4 

2-9-4-0 

44-47 

Nganjoke 

i 

17 

26 

1-9 

55 

17-8 

2-6 

45 

Type  of  B.  micro- 

cephalus, 

Ogowe 

i 

i? 

3i 

2-2 

59 

18-8 

3'2 

45 

Types  of 

longianalis 

2 

15,  J6 

32,33 

2-9,  3'i 

60 

17-5,21-7 

3-i 

46,  47 

Others  from 

Niger  delta 

5 

14-16 

30-32 

1-9-2-75 

54-6i 

18-5-22-3 

2-9-3-5 

45-48 

Tiko,  Cameroon 

i 

16 

30 

1-9 

64 

22-3 

2-9 

47 

Kribi 

i 

17 

28 

2-25 

65 

19-8 

3'9 

47 

Others  from 

Ogowe 

10 

15-17 

27-30 

2-2-2-4 

59-6i 

20-2-23-0 

(2-9) 

45-47 

(in  4) 

3-1-3-8 

«  =  no.  of  specimens,  D  =  dorsal  fin-rays,  A  =  anal  fin-rays,  A/D  =  length  of  anal  base/length  of  dorsal 
base,  sc  =  scales  in  lateral  line,  depth  =  max.  depth  as  %  SL,  c.p.l/d  =  length  of  caudal  peduncle/its  depth, 
vert.  =  vertebrae.  The  Liberian  samples  include  three  syntypes  of  M.  liberiensis. 


TABLE  2 

B.  brachyistius  from  the  extremes  of  its  geographical  distribution.     Data  as  in  Table  i 

Sierra  Leone 

R.  Gambia  (Daget) 

Chiloango 
Matadi 

'Stanley  Falls' 
Niari-Kouilou  (Daget) 

TABLE  3 

Frequencies  of  dorsal  and  anal  fin-ray  counts  in  samples  of  Brienomyrus 

Dorsal  Anal 

13  14  15   16  17  18       24  25  26  27  28  29  30  31  32  33 

Liberia  93  x     4     4     3 

Mungo  462                      1523 

'tnicrocephalus'  and  'longianalis'  1431                                                     2241 

Sierra  Leone  and  Gambia  344                            32141 

Congo,  Chiloango  and  Niari-Kouilou  i           5     7                            751 


n 

D 

A 

A/D 

sc. 

depth 

c.p.l/d 

9 

16-18 

27-31 

1-6-2-1 

(55)  59-63 

18-6-23-4 

2-3-3-3 

i 

18 

30 

1-9 

56 

21-8 

2-85 

6 

15-16 

25-27 

1-8-2-1 

51-56 

17-6-22-0 

2-6-3-0 

i 

13 

26 

1-8 

53 

22-2 

2-4 

i 

15 

25 

1-8 

50 

22-9 

2'55 

5 

15-16 

25 

338  E.   TREWAVAS 

Such  evidence  as  we  have  suggests  variation  both  within  and  between  populations 
no  more  than  may  reasonably  be  expected  in  a  species  occupying  the  lower  reaches 
of  a  series  of  river-systems.  Although  it  is  not  a  typical  inhabitant  of  coastal 
lagoons,  Daget  &  Iltis  (1965)  record  that  it  is  tolerant  of  brackish  water,  so  that  the 
possibility  of  occasional  interchange  between  neighbouring  populations  is  not  ex- 
cluded, perhaps  in  a  geological  rather  than  an  historical  time-scale. 

The  comparative  material  used  for  Tables  1-3  is  that  listed  by  Boulenger  in  vols 
I  and  IV  of  his  Catalogue  under  M.  brachyistius  and  M.  longianalis  with  the  additions 
listed  below.  Some  doubt  attaches  to  the  locality  of  the  specimen  from  'Stanley 
Falls'  since  Poll  &  Gosse  (1963)  did  not  find  B.  brachyistius  in  the  Yangambi  region, 
nor  did  they  include  it  in  their  list  of  species  recorded  from  the  Central  Congo.  The 
collection  by  De  Meuse  in  which  it  was  included  contained  examples  of  several  species 
from  Stanley  Pool  and  of  two  others  from  Stanley  Falls,  Stomatorhinus  microps 
Boulenger  and  Gnathonemus  petersi  Gunther.  Both  the  latter  are  found  throughout 
the  Central  Congo  as  well  as  in  Stanley  Pool  and  the  Lower  Congo,  so  that  they  do 
not  provide  the  confirmation  we  seek  that  De  Meuse's  collection  of  that  date  really 
included  fishes  from  Stanley  Falls. 

Boulenger's  material  included  syntypes  of  M.  liberiensis,  M.  microcephalus  and 
M .  longianalis. 

The  specimen  (BMNH  1970.10.13.1)  registered  as  from  'Lokoja  R.  1967'  may  be 
one  of  the  two  referred  to  by  Reed  (in  Reed  et  al.  1967  :  26  fig.  33)  from  a  swamp  near 
Idah.  He  described  them  as  resembling  M.  brachyistius  but  having  a  shorter  pectoral 
fin  and  a  total  length  of  180  mm.  The  length  of  our  specimen  is  now  only  166  mm. 
In  it  the  right  pectoral  fin  is  reflected  forwards,  but  both  pectorals  when  laid  back 
extend  just  beyond  the  origin  of  the  pelvics,  as  is  usual  in  B.  brachyistius.  The  short 
appearance  of  the  fin  in  Reed's  sketch  may  be  accounted  for  by  the  fact  that  the 
drawing  was  traced  from  a  photograph,  possibly  with  a  foreshortened  fin.  This  fish, 
with  15  dorsal  and  30  anal  rays  and  a  long  slender  caudal  peduncle,  resembles  the 
types  of  M.  longianalis  and  others  from  the  Niger  delta  (Table  3). 

Fowler's  account  of  M.  adustus  (1936)  compares  it  with  M.  batesii  Boulenger,  but 
this  species  has  16  scales  around  the  caudal  peduncle  and  fewer  anal  rays.  Fowler 
did  not  mention  the  peduncular  scales  but  his  drawing  shows  fewer  than  16.  If  we 
assume  that  he  undercounted  the  lateral  line  scales  his  specimens  may  well  belong 
to  B.  brachyistius. 

B.  brachyistius  :  material  in  the  BMNH  additional  to  that  listed  in  Boulenger's 
Catalogue  and  on  p.  336  above. 

BMNH  register  no.     SL  (mm)     Locality  Collector 

1938.12.15.31  102  Akim,  nr  Bunsu,  Ghana  F.  R.  Irvine 

1932.2.27.8  121-5  Onitsha  F.  R.  Irvine 

1970.10.13.1  150-5  'Lokoja  R'  W.  Reed 

1952.4.1.5  89-5  Tiko,  Matute  R.  Cameroun  J.  Deveson 

1950.9.22.1-5  49-68  S.W.  Sierra  Leone  T.  S.  Jones 

1958.9.18.8  107  S.W.  Sierra  Leone  T.  S.  Jones 


FISHES   OF  WEST  CAMEROON  339 

CHARAGIDAE 

Alestes  macrolepidotus  (Val.) 

Brycinus  macrolepidotus  Valenciennes,   1849,  in  Cuvier  &  Valenciennes  vol.   22  :  157  pi.   639 
(Senegal). 

Three  specimens,  42-5-58-5  mm  in  SL  from  R.  Mungo  at  the  bridge  near  Etam, 
coll.  Dr  Disney,  18  March  1970  and  4  Nov.  1970. 

This  widespread  species,  recorded  from  the  Nile,  West  Africa  and  the  Congo,  was 
caught  by  us  only  in  the  part  of  the  Mungo  corresponding  to  its  habitat  in  the 
Yangambi  region,  where  Gosse  (1963)  states  that  it  is  found  near  the  banks  and  along 
sand  banks. 

Our  specimens  were  not  full  grown  and  were  even  more  slender  than  adults  (depth 
26-1-29-2%  SL).  The  gill-rakers  were  rather  fewer  than  the  number  given  by 
Boulenger  for  the  species,  (7-13)  +  (o-i)  +  (15-16)  on  the  first  arch.  The  presence 
of  10  outer  teeth  in  the  upper  jaw,  with  the  second  from  the  symphysis  inset,  out  of 
alignment  with  the  rest,  is  characteristic.  The  dorsal  formula  is  ii-iii  7  or  7  + 1,  the 
anal  iii  11-12.  There  are  23  or  24  scales  in  the  lateral  line,  A\  rows  between  it  and 
the  origin  of  the  dorsal  fin,  i|  from  lateral  line  to  pelvic. 

Colour  notes  made  from  the  smallest  specimen  when  it  had  been  if  hours  in  for- 
malin describe  the  body  as  pale  gold  and  grey,  the  dorsal  fin  with  pink  tinge  and  grey 
leading  edge,  adipose  red,  caudal  lobes  yellow  with  orange  tinge  on  upper  and  lower 
edges  ;  a  diffuse  dark  blotch  at  end  of  caudal  peduncle  continued  as  a  black  streak 
on  middle  caudal  rays.  In  the  preserved  specimens  a  dark  mark  behind  the  pectoral 
girdle  is  more  evident  in  one  specimen  that  was  fixed  in  formalin  than  in  the  others 
where  alcohol  was  the  fixative. 

Alestes  longipinnis 

Brachyalestes  longipinnis  Giinther,  1864,  Cat.  Fish.  vol.  5  :  315  (Sierra  Leone). 

One  specimen  from  the  Mungo  bridge  near  Etam  is  76  mm  in  SL.  It  is  probably 
male,  having  the  longest  dorsal  fin-ray  140%  length  of  head  (near  the  values  obtained 
by  Thys,  1967  :  31).  The  lateral  line  scales  number  31  +  1,  rows  between  lat.  line 
and  dorsal  origin  5|,  between  lat.  line  and  pelvic  2|.  Dorsal  ii  8,  anal  iii  19  +  1. 
Teeth  in  upper  jaw  6  outer,  evenly  aligned,  8  inner;  in  lower  jaw  8  +  2.  Gill- 
rakers  on  first  arch  10  + 1  + 10,  the  epibranchial  ones  very  short. 

No  notes  were  made  on  our  living  specimen,  but  colour  notes  accompanying  one 
collected  in  1952  by  Mr  Deveson  in  R.  Matute  near  Tiko  agree  essentially  with  those 
published  by  Loiselle  (1972)  for  the  species  in  West  Africa,  as  follows  :  eye  brilliant 
gold  (iris  silvery-yellow  with  orange  crescent  on  top  -  Loiselle)  ;  black  mark  on 
caudal  peduncle  and  middle  caudal  rays  with  gold  above  it  (metallic  orange  above  it) ; 
dorsal,  adipose,  caudal  and  anal  fins  with  dashes  of  red  (D,  C  and  A  rosy  violet,  D 
orange  basally).  Loiselle  adds  that  the  long  dorsal  and  anal  filaments  of  mature 
males  are  metallic  white.  From  Thys's  (1967)  account  including  comparisons  with 
specimens  from  the  extremes  of  its  distribution,  the  colour  may  be  more  variable. 
In  particular,  the  prolongations  of  dorsal  and  anal  fins  may  be  reddish. 


340  E.   TREWAVAS 

The  species  inhabits  flowing  water  with  forest  shade  from  Guinea  Republic  to  R. 
Kribi  (excluding  the  Volta  and  the  Dahomey  gap),  as  well  as  similar  habitats  in 
Fernando  Poo. 


CYPRINIDAE 

Labeo  camerunensis  n.  sp. 

PI.  3 

HOLOTYPE.  162  mm  SL  from  R.  Mungo  at  the  bridge  near  Etam,  coll.  Dr  Disney. 
BMNH  1973.5.14.324. 

PARATYPES.  Two  of  SL  167  and  175  mm  from  R.  Wowe  and  two  young  of  43 
and  45-5  mm  from  R.  Mungo  between  Baduma  and  Bolo,  all  coll.  Dr  Disney. 
BMNH  1973.5.14.322-3,  325-6. 

These  fall  into  a  group  of  Labeo  having  the  following  characters  :  eyes  supero- 
lateral,  snout  much  longer  than  postocular  part  of  head,  dorsal  fin  iii-iv  9-10,  its 
longest  ray  not  or  but  little  longer  than  head  and  its  dorsal  edge  concave  ;  16  scales 
around  caudal  peduncle. 

The  branched  dorsal  rays  are  9  in  four  specimens,  10  in  one.  The  snout  is  tuber- 
culate,  without  a  deep  furrow  between  the  ethmoid  and  the  jaw,  projecting  a  short 
way  in  front  of  the  broad  mouth.  A  maxillary  barbel  is  present,  hidden  in  the  folds 
of  the  Up  in  adults  ;  a  short  rostral  barbel  is  present  in  one  of  the  young,  represented 
by  a  stump  or  absent  in  the  others. 

There  are  35  or  36  scales  in  the  lateral  line,  excluding  1-3  on  the  caudal  fin, 
4|-5  rows  above  the  lateral  line  in  front  of  the  dorsal  fin,  3-3!  between  the  lateral 
line  and  the  pelvic. 

Proportions  in  the  three  adults  are  : 

As  %  SL  :  depth  of  body  22-7-23-8  ;  length  of  head  24-7-25-2  ;  length  of  caudal 
peduncle  about  16-17,  its  depth  12-6-13-0  ;  length  of  longest  dorsal  ray  23-0- 
26-2  ;  length  of  pectoral  fin  24-2-25-0. 

As  %  length  of  head  :  diameter  of  eye  20-0-20-7  ;  length  of  snout  55-2-58-5, 
1-65-2-02  times  postocular  part  of  head  including  skinny  edge  of  operculum  ;  inter- 
orbital  width  41-6-46-2. 

Colour  fairly  uniform  on  back  and  sides,  each  flank  scale  in  life  reddish  purple  with 
a  black  base  overlapped  by  the  transparent  posterior  edge  of  the  preceding  scale. 
Lower  side  of  head  and  belly  white  or  pale  grey. 

The  two  young  already  have  the  adult  characters  of  large,  superolateral  eyes 
(25-5,  26-0%  length  of  head),  a  well-developed  snout  with  some  tubercles  and  the  edge 
of  the  dorsal  fin  slightly  concave.  There  is  a  large  round  black  blotch  on  each  side 
at  the  end  of  the  caudal  peduncle  and  a  suggestion  of  a  darker  band  along  the  middle 
of  the  side. 

AFFINITIES.  L.  annectens  Boulenger,  described  from  R.  Kribi  and  neighbouring 
rivers,  has  only  12  scales  around  the  caudal  peduncle  (14  in  one  of  8  specimens 
examined),  but  is  evidently  nearly  related  to  our  species.  L.  batesii  Boulenger, 


FISHES   OF  WEST   CAMEROON  341 

1911  and  1916,  is  known  in  the  adult  state  only  from  the  type,  from  R.  Kribi,  a  fish 
of  SL  169  mm  with  a  broken  neck.  Some  young,  up  to  21-5  mm  SL,  from  R.  Benito 
have  been  attributed  to  it  by  Roman  (1971).  The  drooping  head  of  Boulenger's 
figure  (1916,  fig.  127),  a  restoration,  is  partly  the  result  of  damage,  but  the  high 
curved  back  and  deep  body  (28-4%  SL)  are  natural  and  contrast  with  the  Mungo 
fishes,  whose  dorsal  outline  is  continued  almost  horizontally  into  the  line  of  the  top 
of  the  head  and  snout.  The  caudal  peduncle  of  L.  batesii  is  not  only  deeper  (14-8% 
SL)  and  shorter  (14%  SL),  but  more  compressed  than  in  L.  camerunensis.  The  scale 
numbers  are  only  a  little  higher  than  in  the  Mungo  fish,  those  around  the  caudal 
peduncle  likewise  16,  and  the  length  of  the  longest  dorsal  fin-ray  is  comparable 
(23-6o/0  SL). 

From  the  Sanaga  system  Holly  has  recorded  (ig27a  :  131  ;  ig2jb  :  421  ;  1930) 
L.  greenii  Boulenger  and  Pellegrin  named  L.  chariensis  var.  nunensis  (ig29a). 
Holly  gives  no  details  but  Pellegrin  distinguished  his  'variety'  from  L.  chariensis  by 
the  smaller  and  more  numerous  scales  (1.1.  39,  D  to  1.1.  5^)  and  this  would  place  it  in 
L.  greenii  (1.1.  37-38,  D  to  1.1.  5^-6^).  Both  L.  chariensis  and  L.  greenii  are  further 
distinguished  from  the  Mungo  form  by  the  very  long  last  simple  dorsal  fin-ray 
(37-43%  SL  in  L.  greenii,  about  the  same  in  L.  c.  nunensis}.  These  differences  out- 
weigh a  resemblance  in  the  pigment  pattern  between  the  young  of  the  Mungo  form 
and  L.  greenii,  which  has  a  conspicuous  black  blotch  at  the  end  of  the  caudal  peduncle 
persisting,  though  more  vaguely  delimited,  at  a  standard  length  of  182  mm  (see 
Boulenger's  original  figure,  1902,  pi.  viii  fig.  4).  Pellegrin  mentions  no  pattern 
except  the  usual  countershading,  but  the  syntypes  of  L.  c.  nunensis  were  bigger 
(SL  260  and  330  mm)  than  Boulenger's  specimens  of  L.  greenii. 

L.  mungoensis  is  a  member  of  or  closely  related  to  the  super  species  including  L. 
forskalii,  and  L.  cylindricus,  having  a  lower  number  of  scales  than  these,  but  not  as 
low  as  L.  annectens.  A  dark  lateral  band  is  found  also  in  the  young  of  these  species. 


BARBUS  Cuvier  &  Cloquet,  1816 

I.  Species  with  parallel-striate  scales  and  with  the  last  simple  dorsal  ray  thick 
and  rigid  in  its  proximal  part,  but  not  serrate. 

Our  samples  of  this  group  comprise  75  specimens,  of  which  39  are  young  fry, 
26-56  mm  in  SL.  Good  samples  of  adults  were  taken  in  R.  Nyoke,  Barombi  Mbo 
and  R.  Wowe  ;  R.  Menge  yielded  one  of  92  mm  SL.  In  addition  there  were  14  from 
Lake  Soden  (p.  345). 

These  agree  in  numbers  of  scales  and  fin-rays  and  in  having  two  pairs  of  barbels, 
the  posterior  in  adults  longer  than  the  diameter  of  the  eye  (up  to  twice),  and  in  the 
pharyngeal  dentition,  but  they  fall  into  two  groups  on  the  length  of  the  strengthened 
dorsal  ray  and  its  rigid  part. 

The  ray  is  longer  and  stronger  in  adults  from  Rivers  Wowe  and  Menge  than  in 
those  from  Barombi  Mbo  and  R.  Nyoke.  This  seemed  to  be  the  only  character 
differentiating  them  and  as  it  has  been  found  variable  in  some  other  populations  I 
was  uncertain  of  its  significance  until  I  examined  the  fry.  These  have  longer  dorsal 


342  E.   TREWAVAS 


30 
20 

10 

_i 

oo 

-*  35 


o  o   go      oo»;W  Oo         oo    o      o 

80    100    120    140    160    180   200   220   240   260 


o 
Q 


30 
25 
20 


0       0<b  o< 

O       Ch  O 


25    30    35    40    45    50    55    60    65    70 
SL  mm 

FIG.  2.  Barbus  batesii  (open  circles)  and  B.  mungoensis  (solid  circles).  Length  of  longest 
dorsal  ray  as  %  SL  in  (above)  adults  and  (below)  juveniles.  For  adults  only  the  rigid 
part  of  the  ray  is  measured,  for  juveniles  the  whole  ray.  With  the  adult  B.  batesii  are 
included  the  holotype  and  3  other  Kribi  specimens  ;  the  rest  are  from  Barombi  Mbo  and 
R.  Nyoke.  Triangles  are  ratios  from  Holly's  (1927)  measurements  of  B.  brevispinis. 


and  caudal  fins  associated  with'  longer  barbels,  shorter  fins  with  shorter  barbels. 
The  time  and/or  rate  of  development  of  the  barbels  is  a  feature  that  Barnard 
(1943  :  147)  found  useful  in  differentiating  the  species  of  South  Africa. 

It  happens  that  we  have  a  good  series  from  fry  to  adults  from  only  one  locality, 
R.  Wowe  (Dr  Disney's  collection),  but  this  gives  confidence  in  the  taxonomic  value 
of  the  length  of  the  stiffened  dorsal  ray  at  a  size  when  the  difference  in  barbel-length 
has  been  outgrown. 

Having  separated  the  adults  on  this  basis,  I  was  able  to  find  some  taxonomic 
significance  in  the  degree  of  development  of  the  lips,  a  notoriously  variable  character, 
as  many  authors  from  Gunther  (1868  :  84)  to  Daget  &  Iltis  (1965  :  99)  have  em- 
phasized. It  seems  that  in  the  short-finned  species,  which  I  identify  with  B.  batesii 
Boulenger,  the  lip  may  become  lobed,  but  only  in  large  adults  ;  in  the  long-finned 
species,  to  which  I  have  reluctantly  given  a  new  name,  quite  small  specimens  may 
develop  the  lobe. 

The  two  species  are  contrasted  as  follows  : 
B.  batesii 

Last  simple  ray  of  dorsal  fin  17-24%  SL  in  adults  of  116-314  mm  SL,  shorter 
than  head  ;  its  rigid  part  14-20%  SL.  Lower  lip  with  a  median  pad,  more  or  less 
undercut,  sometimes  produced  as  a  lobe  in  individuals  over  200  mm  SL.  Posterior 
barbel  in  fry  of  33-53  mm  SL  34'5-50-o%  diameter  of  eye. 


FISHES   OF  WEST  CAMEROON 


343 


B.  mungoensis  n.  sp. 

Last  simple  ray  of  dorsal  fin  25-5-30-0%  SL  in  adults  of  114-180  mm  SL,  as  long 
as  or  longer  than  head ;  its  rigid  part  24-5-28-0%  SL,  92-104%  length  of  head. 
Lower  lip  with  an  undercut  or  fringed  median  pad  or  with  a  lobe.  Posterior  barbel 
in  fry  of  25-56  mm  SL  71-100%  diameter  of  eye. 


Barbus  batesii  Boulenger,  1903 

Boulenger,  1903  :  25  pi.  iii  fig.  2  (R.  Kribi)  ;  id.  1911  :  43  fig.  24  ;  Trewavas,  1962  :  152  (Barombi 
Mbo)  ;   Thys  van  dem  Audenaerde,  1967  :  34  figs,  n,  12  (Fernando  Poo). 

In  addition  to  the  material  listed  by  me  in  1962,  we  now  have,  from  our  collections 
of  1970  and  one  made  by  Dr  C.  A.  Wright  in  1963,  the  following  (see  also  p.  345)  : 


18 
17 

2 

6 

4 


SL  (mm)  Locality 

69-5-267  Barombi  Mbo  and  its  feeder  stream 

27~53  R-  Kumba  above  the  falls 

27  and  41  R.  Kumba  at  Buea  Road  Bridge 

28-50  R.  Menge 

155-202  R.  Nyoke 


Proportions,  for  three  size-groups  from  Barombi  Mbo  with  a  separate  column  for 
R.  Nyoke,  are  set  out  in  Table  4  and  proportions  for  fry  up  to  SL  53  mm  are  listed 
in  Table  7  and  Fig.  2. 


TABLE  4 
Proportions  in  Barbus  batesii  in  Barombi  Mbo  and  R.  Nyoke 


SL  (mm) 

N 

Depth  of  body  (%  SL) 

Length  of  head  (%  SL) 

Last  simple  D  ray 

(%  SL) 

(%  length  of  head) 
Rigid  part  of  ray 

(%  SL) 

(%  length  of  head) 
Length  of  pectoral  (%  SL) 
Length  of  anal  (%  SL) 
Length  of  caudal  lobe  (%  SL) 
Snout  (%  head) 
Eye  (%  head) 

Interorbital  width  (%  head) 
Posterior  barbel/eye 


Barombi  Mbo 


Nyoke 


202-267 

116-191 

69-5-78-0 

156-5-202 

4 
24-4-27-8 
23-8-26-6 

ii 
24-8-27-4 
24-5-27-5 

3 

24-8-29-5 
26-8-28-0 

4 
27-8-29-4 
27-7-28-9 

i7«o-20'3 
67-5-85-0 

18-9-24-0 
71-5-89-0 

22-4-28-0 
80-0-100-0 

19-8-21-2 
69-0-77-0 

14-0-16-6 
54-0-70-0 

18-0-20-0 
54-0-74-0 

16-5-22-0 
60-0-80-0 

16-0-17-0 
55-5-59-0 

2I-O-22-2 

20-0-23-0 

22-0-23-0 

22-0-24-5 

19-5-20-5 
24-8-30-4 

33'3-39-Q 
18-6-20-8 
34-8-36 

19-0-23-5 
26-0-30-5 
32-0-39-0 
19-0-30-0 
31-4-36-0 

21-0-21-5 

28-8-31-6 
31-0-34-5 
28-5-32-0 

20-0-24-5 
26-8-28-0 
34-0-39-8 
22-5-23-5 
30-0-34-5 

1-2-1-55 

1-0-1-7 

0-75-1-0 

I-I5-I-5 

344  E-   TREWAVAS 

Scales  24-26  :  4^  (5)  :  2-2^.  5^  between  origin  of  dorsal  and  lateral  line  on  one 
side  of  one  fish.  Dorsal  iii-iv  8-9  in  Barombi  Mbo,  iv  9-10  at  R.  Nyoke.  Gill- 
rakers  (2-4) +  (o-i)  + (9-11)  at  Barombi  Mbo,  (2 -4)  +  !  + (11-13)  at  R.  Nyoke. 

Colour  in  life  brassy  yellow  on  the  flanks,  countershaded  to  dark  green  on  the  back 
and  top  of  the  head  and  white  ventrally ;  anal  fin  colourless,  or  orange  distally 
edged  with  black  ;  base  of  each  scale  marked  with  a  conspicuous  black  crescent. 

For  ecology  see  Green  &  Corbet  in  Trewavas,  Green  &  Corbet,  1972. 

Reasons  for  regarding  B.  linnellii  as  a  synonym  of  B.  batesii  have  been  given  already 
(Trewavas,  1962)  and  comparison  of  additional  material  from  Barombi  Mbo  with  the 
type  of  B.  batesii  and  five  other  Kribi  specimens  confirms  the  synonymy.  Three  of 
the  Kribi  fishes  are  bigger  than  our  biggest ;  in  them  the  positive  allometric  trend  of 
the  snout  and  the  negative  one  of  the  eye  are  continued  to  42%  and  16%  respectively 
of  the  length  of  head.  The  interorbital  width  also  rises  to  39%.  Two,  of  268  and 
314  mm  SL,  have  the  lower  lip  produced  as  a  lobe. 

The  pharyngeal  teeth  of  one  Kribi  specimen  dissected  for  the  purpose  seem  to  be 
in  process  of  replacement  (or  final  shedding  ?  No  new  teeth  were  found  in  the  mem- 
brane), but  teeth  or  seats  on  the  bones  were  present  for  the  usual  formula,  2.3.5-5.3.2 
and  the  shape  was  as  in  the  Barombi  Mbo  specimens.  In  both  this  species  and  B. 
mungoensis  the  anterior  tooth  of  the  row  of  5  is  rather  small,  conical,  the  next  stout 
with  a  point,  the  others  sloping  backwards  with  a  posterior  point  and  an  anterior 
grinding  surface.  In  the  Kribi  specimens  the  posterior  barbel  is  1-5  to  twice  the  dia- 
meter of  the  eye,  gill-rakers  (2-3)  + 1  +  (9-12),  dorsal  iv  9  in  all,  the  last  simple  ray 
69-88-5%  length  of  head,  its  rigid  part  55'5~75'5%.  Scales  26-28  :  4^  :  2-2^. 

The  higher  lateral  line  counts  suggest  a  population  difference  in  this  character,  but 
a  slight  one  (Table  5) .  Thys's  counts  (1967)  from  8  specimens  of  Fernando-Poo  nearly 
cover  both  ranges. 

On  the  basis  of  Holly's  descriptions  (1927, 1929, 1930)  neither  Thys  nor  I  can  recog- 
nize names  given  by  him  to  Barbus  of  Cameroun  as  synonyms  of  B.  batesii.  The 
nearest,  as  Thys  remarks,  is  B.  versluysii  Holly,  1929,  which  has  4-^  scales  from  dorsal 
to  lateral  line  and  fins  of  the  right  proportions  ;  but  Holly  counts  29-30  scales  in  the 
lateral  line  and  his  figure  (1930  pi.  i  fig.  8)  shows  this  count  to  be  made  in  the  same 
way  as  ours  ;  his  description  and  figure  of  the  lips  suggest  greater  development  than 
we  find  in  such  small  specimens  of  our  populations.  Probably  the  posterior  position 
of  the  pelvics  shown  in  the  figure  (Holly,  1930  pi.  i  fig.  8)  is  a  mistake  since  the  de- 
scription records  them  as  under  the  first  rays  of  the  dorsal  (as  in  B.  batesii).  If  this  is 
B.  batesii  it  extends  the  distribution  to  the  Bakoko  highlands. 

The  only  possible  senior  synonym  of  B.  batesii  is  B.  compinei  Sauvage,  1879,  of 
R.  Ogowe,  but  a  decision  on  this  awaits  further  knowledge  of  the  Barbus  population 
of  that  river. 

Blache  et  al.  (1964)  record  three  young  from  the  Chad  basin  (Guelta  de  Tottous) 
as  B.  batesii,  but  from  the  data  given  they  are  more  probably  B.  foureaui  Pellegrin. 

Steindachner  (1914)  recorded  two  species  of  Barbus  from  R.  Ja,  a  secondary 
tributary  of  R.  Congo  rising  near  the  sources  of  the  rivers  of  southern  Cameroun. 
One  of  them,  which  he  identified  as  B.  mawambiensis  Steindachner  (1911),  seems  more 
like  B.  batesii  than  B.  mawambiensis,  the  type  of  which  came  from  Ituri.  The  fauna 


FISHES   OF  WEST   CAMEROON  345 

of  the  Ja  has  much  in  common  with  that  of  the  rivers  of  Cameroun,  arid  it  is  quite 
possible  that  the  distribution  of  B.  batesii  extends  to  it. 

Barbus  batesii  in  Lake  Soden 

Fourteen  specimens,  SL  67-5-219-5  mm,  were  collected  by  Dr  J.  Griffith. 

We  include  them  in  B.  batesii  while  noting  the  following  differences. 

Although  the  scale-counts  for  the  lateral  line  are  the  same  (24  or  25,  27  in  one)  the 
range  between  the  lateral  line  and  the  dorsal  (3|~4|)  is  lower  though  overlapping  ; 
5  of  the  14  specimens  have  3^. 

In  the  size-range  corresponding  to  the  middle  Mbo  column  of  Table  4  (SL  117- 
219-5  mm)  the  length  of  head  is  27-8-30-0%  SL,  longer  than  in  Barombi  Mbo  but 
equal  to  that  of  the  Nyoke  specimens. 

In  the  same  size-range  the  length  of  the  last  simple  dorsal  ray  is  20-5-23-2%  SL, 
thus  within  the  range  for  the  corresponding  Mbo  fishes,  but  the  rigid  part  is  weaker 
and  merges  so  imperceptibly  into  the  flexible  part  that  a  separate  measurement 
cannot  be  made  of  it.  An  attempt  on  some  specimens  gave  11-5-15-0%  SL. 

The  problematic  specimen  recorded  in  1962  from  this  region  is  clearly  conspecific 
with  these.  The  minute  tubercles  on  the  snout  are  present  in  the  new  specimens  but 
also  on  some  from  Barombi  Mbo. 

Barbus  mungoensis  n.  sp. 

HOLOTYPE  AND  PARATYPES.  18  specimens,  40-5-179  mm  in  SL,  from  R.  Wowe, 
tributary  of  R.  Mungo,  coll.  Dr  Disney  in  October,  1970.  BMNH  1973.5.14.163 
(holotype,  179  mm),  and  BMNH  1973.5.14.164-182  (paratypes). 

OTHER  MATERIAL  EXAMINED.  5,  30-38  mm  SL,  R.  Blackwater,  i,  92  mm  SL, 
R.  Menge,  and  2,  26  and  43  mm  SL  from  R.  Mungo  between  Baduma  and  Bolo,  all 
coll.  Dr  Disney. 

Scales  24-26  (28)  :  4!  (5^)  :  2.  Dorsal  iii-iv  8-9,  usually  iv  9.  Gill-rakers  on 
first  arch  (2-5)  +  (o-i)  +  (9-12).  Pharyngeal  teeth  2.3.5-5.3.2. 

Proportions  for  three  size-groups  are  set  out  in  Table  5.  The  first  and  second 
columns  correspond  in  size  roughly  to  the  second  and  third  in  Table  4  (B.  batesii), 
the  third  is  a  younger  size-group.  Some  features  of  the  fry  are  compared  with  young 
B.  batesii  in  Table  7  and  Fig.  2.  In  the  26  mm  fish  from  the  Mungo  the  posterior 
barbel  is  already  0-8  diameter  of  eye  and  the  upper  caudal  lobe  31%  SL,  but  at  this 
stage  the  last  simple  dorsal  ray  is  no  thicker  than  the  following  ray. 

A  specimen  of  92-5  mm  SL  from  R.  Menge  agrees  better  with  this  species  than  with 
B.  batesii.  It  has  a  very  long  snout  (42%  length  of  head)  and  flexible,  lobed  lips. 
The  flexible  part  of  the  dorsal  spine  is  broken  off  and  the  rigid  stump  is  only  22-4% 
SL,  but  it  is  thicker  than  in  any  B.  batesii  of  comparable  size.  The  posterior  barbel 
is  as  long  as  the  eye  and  the  upper  caudal  lobe  is  32%  SL.  It  agrees  in  number  of 
scales  (24  :  4!  :  2)  and  gill-rakers  (3  +  1  +  11)  with  both  the  Mungo  B.  batesii  and 
B.  mungoensis.  The  few  fry  from  the  Menge  are  probably  B.  batesii  and  the  two 
species  may  exist  together  in  this  tributary. 


346 


E.   TREWAVAS 


TABLE  5 

Barbus  mungoensis  from  R.  Wowe 


SL  (mm) 

N 

Depth  of  body  (%  SL) 

Length  of  head  (%  SL) 

Last  simple  D  ray 

(%  SL) 

(%  length  of  head) 
Rigid  part  of  ray 

(%  SL) 

(%  length  of  head) 
Length  of  pectoral  (%  SL) 
Length  of  anal  (%  SL) 
Length  of  caudal  lobe  (%  SL) 
Snout  (%  head) 
Eye  (%  head) 

Interorbital  width  (%  head) 
Posterior  barbel/eye 


114-5-179 

3 

28-29 
25-4-26-8 

25-5-30-0 

IOO-III 

24-5-28 

92-104 
21-0-23-8 

19-22 
27-3-3I-4 
35-8-37-6 
24-8-28-4 

31-34 
I-3-I-5 


76-79-5 
4 

27-5-30-5 
27-5-3I-6 

30-0-34-0 
106-120 

26-6-30-2 
84-108 

22-25 

23-24 
31-2-33-8 
29-39-5 
30-32-5 
28-31-4 
o-86-i-i 


40-5-69-5 
ii 

28-31 
29-3I-5 

30-0-34-0 
91-114 

26-30 

80-100 

22-5-23-5 

3I-5-36 
29-36 
0-8-1-2 


TABLE  6 

Scales  in  the  lateral  line  in  Barbus  batesii  and  B.  mungoensis 
24  25  26  27  { 


B.  batesii 

R.  Kribi  i 

R.  Nyoke  3  I 

Barombi-Mbo  '2  9  7 

R.  Kumba  144 

B.  mungoensis 

R.  Wowe  365 

R.  Blackwater  221 


TABLE  7 

Contrasts  in  length  of  fins  and  barbels  in  fry  of  Barbus  batesii  and  B.  mungoensis 


SL  (mm) 

D  ray  (%  SL) 

C  lobe  (%  SL) 

Pectoral  (%  SL) 

Posterior  barbel  (%  eye) 


B.  batesii  B.  mungoensis 
(R.  Kumba)    (Wowe,  Blackwater,  Mungo) 

27-53  26-56 

21-26  29-35 

27-32  31-36 

17-21  21-23-5 

25-50  71-100 


Life  colours  are  not  recorded.  The  fry  sometimes  have  a  dark  spot  at  the  caudal 
base  and  there  may  be  irregular  smudges  on  the  body  and  caudal  lobes. 

RELATIONSHIPS.  Although  this  population  does  not  correspond  exactly  with 
descriptions  of  others  it  appears  to  be  one  of  a  series  beginning  with  B.  habereri 
(R.  Ja)  and  continuing  with  B.  foureaui  into  the  Chad,  Niger  and  Volta  basins. 


FISHES   OF  WEST   CAMEROON  347 

B.  habereri  Steindachner  (1914  :  24  fig.  4  pi.  iii  fig.  i)  agrees  with  it  in  scale-formula 
(25  :  4^  :  2|)  and  the  well-developed  lips,  but  has  only  one  pair  of  barbels.  Dr 
Kahsbauer  has  recently  kindly  confirmed  this  for  me  and  gives  the  gill-raker  count 
as  4  + 1 +  n.  According  to  Steindachner's  figure  the  rigid  dorsal  ray  is  34-6%  SL. 

B.  brevispinis  Holly,  1927  (Rivers  Lokundje  and  Sanaga,  Cameroun)  in  spite  of  its 
name  has  a  dorsal  spine  as  long  as  in  B.  mungoensis,  but  has  30  scales  in  the  lateral 
line  series  and  5|  between  it  and  the  origin  of  the  dorsal. 

Daget  (1954  :  188),  and  following  him  Blache  et  al.  (1964)  and  Daget  &  Iltis  (1965), 
have  used  the  name  B.  occidentalis  Boulenger  for  the  Chad  and  western  populations, 
but  the  holotype  of  B.  occidentalis  has  a  short  dorsal  spine,  its  rigid  part  only  three- 
quarters  length  of  head.  Until  the  Ogun  population  is  studied  I  think  it  safer  to  use 
B.  foureaui  Pellegrin,  igigb  (syn.  B.  seguensis  Pellegrin,  1925)  for  these  populations. 
They  differ  from  the  Mungo  population  in  having  usually  5|  scales  between  the  dorsal 
and  the  lateral  line,  12-16  gill-rakers  on  the  lower  part  of  the  first  arch  and  the  lips 
more  constantly  lobed. 

B.  lancrenonensis  Blache  &  Miton,  1960  (see  also  Blache  et  al.  1964  :  126,  fig.  60) 
from  R.  Ngou,  affluent  of  R.  Mbere,  Logone  system,  was  described  from  three  tiny 
specimens  14-7-33-5  mm  long.  The  authors  thought  that  it  belonged  to  the  section 
of  Barbus  containing  small  species  with  no  enlarged  dorsal  ray  and  with  radiately 
striate  scales,  but  several  features  point  to  its  being  the  young  of  B.  foureaui.  The 
radii  of  the  scales  were  described  as  'relativement  nombreuses  et  faiblement  conver- 
gentes'  ;  dorsal  iv  9  ;  gill-rakers  10-11  on  lower  part  of  arch  ;  a  few  lines  of  'pores' 
on  the  head  (as  I  have  noted  too  in  fry  of  the  Mungo  system)  ;  posterior  barbel 
o-6-i-o  diameter  of  eye.  It  is  the  similarity  between  these  and  our  young  B. 
mungoensis  that  drew  my  attention  to  them  and  leads  me  to  suggest  their  identity 
with  B.  foureaui. 


II.  There  are  three  species  of  Barbus  with  radiating  striae  on  the  scales  and  with 
the  last  simple  dorsal  ray  flexible,  not  serrate. 

Barbus  (Enteromius)  callipterus  Boulenger,  1907 

Boulenger,  igoya  :  486  (R.  Kribi  at  Akok)  ;   id.  1911  :  167  fig.  145  ;   Thys  van  den  Audenaerde, 
1967  :  48  fig.  15  (Fernando  Poo). 

For  use  of  the  subgenus  Enteromius  Cope,  1869,  see  Greenwood  (1970),  who  shows 
that  the  name  should  replace  Beirabarbus. 

This  species  was  caught  in  R.  Mungo  between  Baduma  and  Bolo,  in  the  Mungo 
tributaries  Wowe  and  Menge  and  was  the  only  small  Barbus  caught  by  us  in  the 
Kotto  area,  being  taken  in  the  lake  itself,  in  Tung  Nsuia,  a  stream  entering  it  from 
the  crater  rim,  and  in  the  outflowing  stream  Nganjoke. 

All  Kotto  specimens  had  the  characteristic  black  patch  on  the  dorsal  fin  leaving 
the  tip  of  the  fin  pigment-free.  In  life  the  proximal  part  of  this  fin  varies  from  pale 
yellow  or  salmon  pink  to  orange  and  the  caudal  fin  from  pale  yellow  to  orange-red. 
In  a  well-coloured  fish  the  top  of  the  iris  is  red.  In  all,  the  head  is  brassy  yellow. 


348  E.   TREWAVAS 

Already  at  14-18  mm  SL  these  colours  are  developed  though  still  at  the  pale  end 
of  the  range. 

In  samples  from  the  Mungo  and  its  tributaries,  where  it  was  abundant,  the  black 
pigment  on  the  dorsal  fin  is  much  less  intense,  often  faint,  less  sharply  demarcated 
than  in  the  Kotto  samples  or  the  types,  and  may  extend  nearly  to  the  end  of  the 
first  three  branched  rays. 

No  significant  differences  in  numbers  of  scales,  fin  rays  and  gill  rakers  could  be 
found  between  the  two  populations  and  the  pharyngeal  teeth  are  alike. 

Comparison  in  proportions  between  Kotto  and  Mungo  samples  is  blurred  because 
in  the  preserved  material  all  Kotto  specimens  (except  the  two  juveniles)  are  bigger 
than  all  Mungo  specimens.  The  four  syntypes  of  B.  callipterus  are  also  smaller 
than  the  Kotto  specimens,  so  that  when,  as  with  length  of  head  and  diameter  of  eye, 
the  Mungo  specimens  are  more  like  the  types  the  difference  between  both  and  the 
Kotto  fishes  can  be  ascribed  to  allometry.  Only  in  the  height  of  the  dorsal  fin  is  the 
Mungo  sample  possibly  peculiar.  This  ratio  is  : 

in  8  specimens  of  64-7-73-5  mm